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Full text of "Bulletin of the Natural History Museum Zoology"

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ISSN 0968-0470 



Bulletin of 

The Natural History 

Museum 



THE NATURAL 
HISTORY MUSEUM 

-9 JUL 1993 

PhiESENTfcD 
ZOOLOGY LIBRARY 



Zoology Series 





THE 

NATURAL 
HISTORY 
MUSEUM 



VOLUME 59 NUMBER 1 24 JUNE 1993 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum 
(Natural History)), instituted in 1949, is issued in four scientific series, Botany, 
Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Zoology Series is edited in the Museum's Department of Zoology 
Keeper of Zoology: Dr C.R. Curds 

Editor of Bulletin: Dr N.R. Merrett 

Assistant Editor: Dr B.T. Clarke 



Papers in the Bulletin are primarily the results of research carried out on the unique and ever- 
growing collections of the Museum, both by the scientific staff and by specialists from elsewhere 
who make use of the Museum's resources. Many of the papers are works of reference that will 
remain indispensable for years to come. All papers submitted for publication are subjected to 
external peer review for acceptance. 

A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. 
Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers 
and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and 
enquiries should be sent to: 

Intercept Ltd. 
P.O. Box 716 
Andover 
Hampshire SP10 1YG 

Telephone: (0264) 334748 
Fax: (0264) 334058 



Wo rid List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) 
© The Natural History Museum, 1993 



Zoology Series 
ISSN 0968 - 0470 Vol. 59, No. 1, pp. 1-96 

The Natural History Museum 

Cromwell Road 

London SW7 5BD Issued 24 June 1993 

Typeset by Ann Buchan (Typesetters), Middlesex 
Printed in Great Britain at The Alden Press, Oxford 



Bull. nat. Hist. Mus. (Zool.) 59(1): 1-9 



Issued 24 June 1993 



A new snake from St Lucia, West Indies 



THE N ;L 

HISTC UM 



GARTH UNDERWOOD 

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5 Bl 



"9 JUL 1993 

Ph^b'ENTED 

ZOOLOGY LiBRA RV 



Synopsis. The Clelia clelia reported from Dominica and St Lucia are reinvestigated. The specimens concerned are 
recognised as a new species, Clelia errabunda. It is derived in relation to the mainland species C. bicolor and C. 
rustica and is the primitive sister species of the mainland C. clelia, C equatoriana and C. scytalina. As such it is 
interpreted as an island relict. Records from Dominica and Guyana are rejected, the St Lucia records are confirmed. 



INTRODUCTION 



Nearly a hundred years ago Boulenger (1896) reported the 
South American snake Oxyrhopus clelia, now known as 
Clelia clelia, from the Windward Islands of Dominica and St 
Lucia, as well as from mainland localities. Ever since then 
this species has been listed for these two islands, either in the 
genus Clelia (Barbour, 1930; Schwartz & Henderson, 1988) 
or Pseudoboa (Barbour, 1935, 1937). There are other records 
of this snake for St Lucia but this remains the only record for 
Dominica. 

Boulenger (1896) distinguished two varieties of "Oxyrho- 
pus clelia': A, with 19 scale-rows at mid-body, and B, with 17 
scale-rows. Under variety 'B' Boulenger lists: two specimens 
from 'The city of Mexico', three specimens from 'W. Ecua- 
dor' and one specimen each from 'Demerara', 'Dominica' 
and 'St Lucia'. 

Bailey (1970, in Peters & Orejas-Miranda) gives a key to 
the mainland species of Clelia. The city of Mexico specimens 
key out as Clelia scytalina, the Ecuador specimens as C. 
equatoriana. The remaining three specimens reach the 
scytalina-equatoriana couplet of the key but are not clearly 
assignable to either mainland species. Boulenger's variety 'A' 
specimens key out as C.c. clelia and C.c.plumbea, the two 
subspecies recognised by Bailey. Clelia was later reported 
from the island of Grenada. It was described by Greer (1965) 
as Clelia clelia groomei; Bailey places this in the synonymy of 
C.c. clelia. 

Further examination of these remaining three specimens 
shows that they represent an unrecognised species. In the 
hope of determining the affinities of the new form the 
specimens of Clelia in the collection of The Natural History 
Museum, London were examined. A Paris Museum specimen 
from St Lucia was also examined. The seven species recogn- 
ised by Bailey are represented. It should be noted that 
Scrocchi and Vihas (1990) have now sunk occipitolutea in the 
synonymy of clelia. 



MATERIALS AND METHODS 



Each specimen was examined in respect of external features, 
anterior viscera, maxillary teeth and, in the case of males, 
hemipenis. For a representative of each species dissections 
were made to show the superficial jaw muscles, ligaments and 



labial glands and the upper and lower jaws. 

The ventral scale count is according to Dowling (1951). 
The notation for the scale-rows on the trunk allows a detailed 
record in a single line of type using characters on a word 
processor keyboard. An oblique transverse scale-row is iden- 
tified by the ventral scale from which it passes backwards, as 
shown by M.A. Smith (1943, fig. 10). Where there is a 
scale-row fusion the upper of the two merged rows is indi- 
cated. For example male specimen BMNH 89.4.8.2 is 
recorded as: 

V 210 >>5> 17 (36;5</33;5<) 19 (145;4>/142;4>) 17 

Ventral scale count 210. Three scale-row fusions from the 
back of the head onto the neck, the third recognisable as due 
to the fusion of row 5 with the row below. Seventeen rows on 
the neck through oblique row 36 left/33 right; row 5 splits 
giving rise to 19 rows through oblique row 145 left/142 right; 
row 4 merges with row below giving 17 rows. Most specimens 
were recorded down the left-hand side only (Table 1, see 
Appendix). 

The subcaudal count starts with the first scale on the left 
side which makes full contact with a scale obliquely forward 
and opposite. The dorsal scale-rows on the tail are somewhat 
irregular around the base but settle down to even row 
stretches: 8, 6, 4, 2. The fusions are rarely symmetrical so that 
there are short odd number transitions between the even 
stretches. Each transition is included in the preceding, 
higher, even row stretch. The transverse rows are identified 
by the subcaudal scale-row pairs from which they arise 
obliquely. The tail of the above specimen was recorded as: 

79 prs, >8 (10) 8 (25) 6 (45) 4 (73) 2 

More than eight longitudinal rows through oblique row ten, 
eight (and seven) through row 25, six (and five) through row 
45, four (and three) through 73 and two rows to the terminal 
scute. From this record the lengths of the five stretches are 
calculated as the basis of comparison of the individual speci- 
mens. The above becomes: 

C79 >8:10 8:15 6:20 4:28 2:6 

This feature shows marked sexual dimorphism. 

For selected specimens scales were mounted on slides to be 
examined for the presence of pits and tubercles (Underwood, 
1963). The wet scales are laid on a dried film of polyvinyl 
alcohol lactophenol mounting medium. When dry the slide is 
then covered using Canada balsam. For each specimen the 



G. UNDERWOOD 



scales mounted are the frontal, a parietal and a vertical series 
of scales at about mid-trunk from row one to the vertebral. 

The immunological studies of Cadle (1984) suggest that 
species of Pseudoboa are sister to Clelia; his sample included 
the type species of each genus. He also found that Oxyrhopus 
fitzingeri is closer to Clelia plus Pseudoboa than it is to other 
species of Oxyrhopus. 'Oxyrhopus' fitzingeri is therefore 
included as part of the outgroup. In the absence of an 
analysis, the majority outgroup condition is taken to be 
primitive for the ingroup. 

The four species of Pseudoboa recognised by Bailey share 
the special feature of single subcaudals. No evident special 
feature unites the species of Clelia. On further study the 
boundary between the two genera may be redrawn, or 
abolished so that the assignment of the species discussed here 
may be changed. 

The loreal is rather variable. It may be about as large as the 
preocular, in which case it meets supralabials two and three. 
It may be smaller and may meet only supralabial two. It may 
be absent allowing the prefrontals to meet the supralabials, or 
the nasal to meet the preocular (as a unilateral variant, C. 
clelia BMNH 89.4.8.2). A large loreal is assumed to be 
primitive. 

Boulenger (1896) distinguishes between those Oxyrhopus 
(s.l.) in which the preocular reaches the upper surface of the 
head and those in which it does not. the distinction appears to 
be real but it can be influenced by the condition of the 
specimen and by the angle of view. The supraocular scale 
meets the preocular and often, also, the prefrontal scale. The 
lengths of the supraocular-preocular and the supraocular- 
prefrontal sutures are compared. The prefrontal suture may 
range from two or three times the length of the preocular 
suture, as in Clelia rustica, to absent, as in Oxyrhopus petola 
in which the preocular meets the frontal. 

There are one preocular and two postoculars. The tempo- 
rals are 2:3. The upper anterior temporal always meets 
postoculars; the lower may meet them (represented by 
+/+2:3) or may not meet them (+/-2:3). In every species 
represented by more than two specimens this contact is 
variable. Where the temporal does not make contact the 
labial scale is higher than its neighbours. However, rustica is 
nearly constant, on only one side of one of nine specimens 
does the lower temporal make contact (BMNH 95.9.17.21). 
The outgroup is also variable so polarity is uncertain. In one 
rustica there is no suture between prefrontal and preocular 
(BMNH 86.1.19.21). 

There may be three or two anterior supralabials. As three 
is the commonest number in the outgroup it is assumed to be 
primitive. There are two supralabials meeting the eye and 
three postocular supralabials. Anterior and posterior infrala- 
bials are distinguished. The last anterior infralabial is pentag- 
onal; from it starts the posterior row and a mesial row. The 
anterior infralabials are usually five. Four and six occur as 
unilateral variants; one rustica (BMNH 1909.11.2.16) has 
four symmetrically. The posterior infralabials range from 
four to two. As four is the commonest number in the 
outgroup this is assumed to be primitive. 

The genials range from posterior about as large as the 
anterior to somewhat smaller. This does not appear to be a 
taxonomically useful feature. 

There are always two, sometimes three, scale row fusions 
from the back of the head onto the neck. The rows on the 
neck may be 19 or 17. As a majority of the outgroup have 19 
this is inferred primitive. In those with 19 rows this number 



continues through mid-body and then reduces by fusion of 
row four or five with the row below to 17 rows. The vertebral 
or paravertebral rows were not seen to be involved in 
scale-row changes. Those with 17 neck rows may continue 
without change to the end of the trunk. In some specimens, 
however, a lateral row splits on the posterior neck to give rise 
to 19 rows through mid-body, then reducing to 17 rows. Such 
a minimum on the neck rising to a maximum around mid- 
body and falling towards the vent is widespread in henophid- 
ian grade snakes (Underwood and Stimson, 1990). However, 
this condition is found in none of the outgroup so it is here 
assumed to be a derived, pseudoprimitive feature. The scale- 
row pattern is scored as 19:19:17 (inferred primitive), 
17:19:17 or 17:17:17. 

In some specimens the vertebral scale-row is undifferenti- 
ated, but in many it is wider than the paravertebral rows. This 
modification is a little more pronounced towards the poste- 
rior trunk. As most of the outgroup have undifferentiated 
vertebral scales this is assumed to be primitive. 

These snakes usually have both scale pits and tubercles 
(Underwood, 1963). The pits are confined to the apices of the 
scales of the trunk and tail, where they usually occur in 
pairs. Tubercles may be numerous on the head scales. On the 
trunk they are rather irregularly distributed around the centre 
of each scale; numbers range from five to zero (Fig. 2). Both 
pits and tubercles tend to be better developed on the upper 
scale-rows. Where the vertebral scale-row is enlarged 
they may be reduced or missing. Pits and tubercles are 
found in the outgroup and their presence is inferred primi- 
tive. 

The anterior viscera were examined and their positions 
recorded in ventral scale units. This has the advantage that 
juveniles and adults can be compared. Organ positions deter- 
mined by measurement are subject to allometric change 
(Thorpe, 1975). The features recorded are: tip of the hyoid, 
tip of the ventricle, anterior end of the liver and the end of 
the tracheal cartilages. In all the entry of the trachea into the 
right lung is terminal. This is a derived condition which 
Wallach (personal communication) reports to be general in 
Xenodontine snakes. In all there is some extension of the 
vascularisation of the lung into the roof of the trachea, but it 
does not extend far forwards of the heart. The left lung may 
be present but small, up to about one or two scale-units long 
and vascular; it may be a non-vascular vestige or it may be 
absent. Presence is inferred primitive. 

The trachea may terminate no more than three or four 
scale-units beyond the tip of the heart. It may extend to 
overlap the liver, in some cases extending the full length of 
the vascular portion of the lung to reach the terminal air-sac. 
A short trachea is inferred primitive. 

Counts are made of the teeth, and empty tooth places, of 
the left maxilla. The solid teeth show a moderate increase in 
size from front to rear. They are followed by an interval and 
two obliquely placed teeth with anterior grooves (Fig. 3). 
Polarity is not inferred. In one juvenile specimen (clelia, 
BMNH 1933.6.24.102) no grooves could be seen, even when 
the maxilla was removed and dried. The anterior tooth count 
is recorded. For sample specimens counts were made of the 
teeth on the dentary, palatine and pterygoid bones. They all 
have a full length choanal process of the palatine bone with a 
broad base, about half the length of the palatine, which 
sweeps backwards into a process which overlaps the ptery- 
goid bone by two to three teeth (Fig. 3). The maxillary 
process of the palatine is turned backwards; it bears a 



A NEW SNAKE FROM ST LUCIA 



foramen for the maxillary nerve which emerges on the 
underside anteriorly. 

For sample specimens the skin on the side of the neck was 
turned forwards to expose the superficial jaw muscles and 
ligaments and the labial glands. The most superficial muscle, 
which is easily damaged, is the constrictor colli (Haas, 1973). 
In the outgroup this is a thin sheet of muscle which passes 
from about the level of the head of the quadrate backwards 
and downwards over the jaw articulation to insert on the skin 
of the throat. In the species of Clelia examined the muscle 
follows a similar course, the anterior portion has a diffuse 
origin on the surface of the adductor externus profundus 
muscle. The posterior portion arises on the head of the 
quadrate; this is inferred derived (Fig. 4). 

The cervico-mandibularis muscle arises from the back of 
the neck and passes downwards and forwards to insert on the 
articular head of the quadrate (Fig. 4). This appears to be a 
primitive condition. From the articular head of the quadrate 
arises a ligament which passes forwards and divides. The 
lateral, labial portion inserts on the posterior supralabial 
scales and onto the capsule of Duvernoy's gland. It peels off 
the supralabial scales rather more easily than is usual in 
snakes; this is thought to be derived. The mesial, maxillary 
ligament passes forwards to insert on the posterior lateral 
corner of the maxilla. 



undifferentiated vertebral scale row, no left lung and a short 
trachea extending no more than five ventral scale units 
beyond the tip of the ventricle. Distinguished from rustica 
and bicolor by absence of a left lung. Distinguished from 
clelia, equatoriana and scytalina by the undifferentiated verte- 
bral scale-row and short trachea. Further distinguished from 
clelia by 17 scale-rows from neck to vent. 

Holotype: BMNH 89.8.14.25, male, St Lucia, West Indies, 
collected by G.A. Ramage, presented by West Indies Explo- 
ration Committee; snout-vent c.112 cms, tail 32 cms with 
extreme tip missing. 

Paratype: MNHP 7598, male, St Lucia; snout-vent 
c.116 cms, tail 29+ cms with tip missing. 

Referred specimens: BMNH 89.8.14.12, female, 'Domin- 
ica', West Indies, collected by G.A. Ramage, presented by 
West Indies Exploration Committee, snout-vent 138 cms, tail 
27.6 cms. 

BMNH 1988.717, female, 'Demerara', presented by Capt 
E. Sabine, R.E., snout-vent 117 cms, much of the tail is 
missing. 

The other species of Clelia are widely distributed on the 
South and Central American mainland and a few offshore 
islands (Bailey, 1970). The name is taken from the Latin 
errabundus = wandering, in reference to the occurence of 
the new form well outside the range of its mainland relatives. 





Fig. 1. Clelia errabunda sp. nov., type BMNH 89.8.14.25. A, lateral view of head (reversed on account of distortion of right side of head); 
B, dorsal view of head. 



Mucous supralabial glands extend along the margin of the 
upper lip from the corner of the mouth to the snout. There 
are similar glands along the margin of the lower jaw. Mesial 
to the three posterior supralabial scales lies the Duvernoy's 
(venom) gland (Fig. 4). 

At the level of the corner of the mouth, mesial to the 
maxillary ligament, is the organ termed anterior temporal 
gland by Smith & Bellairs (1947) and rictal gland by McDow- 
ell (1986). It is found in all of the species examined; it is 
usually visible mesial to the posterior end of the Duvernoy's 
gland (Fig. 4). 



RESULTS 

Clelia errabunda sp. nov. 

Diagnosis. A species of Clelia with uniform dark grey adult 
colouration of the upperside of the head, trunk and tail, 
extending to the lateral margins of the ventral scales, an 



The type has two preocular scales on one side, seen in no 
other specimen of Clelia. All have two anterior and three 
posterior temporals. The lower anterior temporal scale meets 
a postocular in the type only. The 'Demerara' specimen has 
three anterior supralabial scales on the left-hand side, seen 
elsewhere only in C. bicolor. All have five anterior infralabial 
scales and three posterior. The anterior genials are little, if at 
all, larger than the posterior. The four specimens have 
tubercles but no pits on the head, as in other Clelia and 
Pseudoboa. The number of frontal and parietal tubercles is 
high. Most of the trunk scales bear paired apical pits, as is 
usual in Clelia, and a moderate number of tubercles (Table 2, 
see Appendix). 

The island specimens have 14 anterior maxillary teeth (on 
the left), which is higher than for clelia and equatoriana 
(Table 3, see Appendix); the 'Demerara' specimen has 13/12. 
The ventral scale counts are high for Clelia, but not extreme. 
The subcaudal scales are entirely paired, save that the last 
one is single in the 'Dominica' specimen. Apart from the 
difference of sex the three island specimens are very similar. 



G. UNDERWOOD 




Fig. 2. Clelia errabunda sp. nov., mounted scales to show distribution of tubercles and pits. a,b, parietal and frontal of BMNH 89.8.14.12; 
c,d, frontal and parietal of BMNH 89.8.14.25 (type); e, mid-trunk scales of BMNH 89.8.14.12, from rows: 4, 5, 6, 7, paravertebral and 
vertebral. The lower rows lack scale-organs. 



I I. 




pp / cp 




Fig. 3. Clelia errabunda, BMNH 85.8.14.12. i, mesial view of lower law: a = angular, d = dentary, s = splenial; ii, lateral view of left 
maxilla (reversed); iii, ventral view of left upper jawbones: e = ectopterygoid, f = fangs with grooves, m = maxilla, pa = palatine, 
pt = pterygoid; iv, mesial view of left palatine-pterygoid articulation: cp = choanal process, pp = posterior process of palatine. 



The 'Dominica' female has 234 ventrals and 71 subcaudals: 
total 305. The type male has the extreme tip of the tail 
missing, judged to be not more than two pairs of subcaudals. 
It has 221 ventrals and 82 + (?)2 subcaudals: total 
303 + (?)2. The paratype male has 224 ventrals and 75 + 
subcaudals. The 'Demerara' female has 230.5 ventrals and 
only 36 remaining pairs of subcaudals. 

The 'Dominica' specimen has a Duvernoy's (venom) gland 
from behind the eye to the corner of the mouth; it is as high as 
the supralabial scales plus the lower temporal scales. The 
hemipenes of the two males are 18 subcaudal scale units long, 
there are prominent lobes on a shaft 13 units long. The sulcus 
spermaticus forks on the shaft at scale six (type) or seven 
(paratype). Proximally on the shaft there are very fine spines 



and, in the retracted organ, longitudinal folds. From scale 
nine to the cleft there are large spines, about 26 in the type 
and 38 in the paratype; these are high counts for Clelia. At 
the base of each lobe there is a large spine, as is usual in 
Clelia. the branch sulcus passes down the middle of an area of 
large calyces with a clear margin (a capitulum). 

Inspection of The Natural History Museum register raises a 
doubt about the provenance of the Dominica specimen. G.A. 
Ramage brought back a collection of herpetological speci- 
mens from Dominica and St Lucia. These were registered in 
1889. They are entered in Boulenger's hand. The register 
starts (with present identifications substituted): 

89.8.14. 1-8 Typhlops dominicana Dominica, June '89 



A NEW SNAKE FROM ST LUCIA 



9-11 


Alsophis antillensis 
sibonius 


Dominica, June '89 


12 


Clelia errabunda 


Dominica, June '89 


13 


Bothrops caribbaeus 


Dominica, June '89 


14 


Thecadactylus 
rapicauda 


St Lucia, April '89 



Cm -. 



There follow another eight species attributed to St Lucia, 
including the St Lucia endemics Hyla rubra, Sphaerodactylus 
microlepis, Anolis luciae and Liophis ornatus. these St Lucia 
attributions are not therefore in question. Boulenger's St 
Lucia entries are interrupted by four fish entries in a different 
hand. 

The Typhlops and Alsophis are forms endemic to Domin- 
ica. What, however, attracts attention is the record of Both- 
rops from Dominica. We may be sure that if a pit-viper were 
living on this island there would have been further reports 
since 1889. It is clear that the Bothrops was mistakenly 
attributed to Dominica. The specimen of Clelia appears to be 
the only documented record of the genus from Dominica. Is 
this attribution to Dominica also a mistake? On the other 
hand there are several further specimens of Clelia from St 
Lucia in the Museum of Comparative Zoology. 

In addition to Alsophis antillensis, there is no more than 
hearsay evidence of a second species of black snake on 
Dominica. Bullock & Evans (1988) list Clelia clelia on 
Dominica as 'Tete-chyen nwe'. Dr Bullock writes that he has 
not seen Clelia but he has reports from informants whom he 
regards as reliable. Dr R. Thorpe, Miss A. Malhotra and Mr 
M. Day have come across no evidence of Clelia on the island. 
On Dominica it would be distinguished from Alsophis by the 
uniform black dorsal colouration and by 17 dorsal scale rows 
on the anterior trunk. The Alsophis has some irregular yellow 
markings and 19 scale rows anteriorly. Unless and until there 
is clear evidence of the occurrence of Clelia on Dominica it 
should be dropped from the island list. Barbour (1930) says of 
'Clelia clelia' that "This species is surely extinct on St 
Lucia. . .". Long (1974) writes that "the cribo no longer 
exists in St Lucia. . .". Dr D. Corke also reports that he has 
found no trace of the survival of Clelia on St Lucia. 

Even greater doubts arise about the provenance of the 
'Demerara' specimen of errabunda. The specimen had no 
original registration number; the Museum register starts in 
1837. A search of the early entries shows no record of 
specimens from E. Sabine. The Museum archivist reports 
that the trustees' minutes record donations from Capt Sabine 
between 1818 and 1824 with, however, no indication of any 
from the Caribbean. 

The collection has other snakes from 'Edw. Sabine'. There 
is a male Xenodon merremi, a species widely distributed in 
South America and known from Guyana on the basis of other 
specimens. There are a male and a female of Oxyrhopus 
trigeminus, not otherwise known from this part of South 
America. 

There are also two lots of Bothrops. A female from 'Capt 
Sabine, Berbice' and two females and a male from 'Col 
Sabine, Demerara'. These were compared with specimens of 
B. atrox and B. brazili from Guyana and with B. caribbaeus 
from St Lucia. With ventral and mid-body counts of: M 
201:26, F 205:27, F 206:29 and F 210:29 they fall within the 
range of counts reported by Lazell (1964) for caribbaeus. The 
postocular stripe passes across the last supralabial scale dorsal 
to the corner of the mouth as in caribbaeus. The ventral scales 
are laterally peppered with dark spots, as in caribbaeus and 




Fig. 4. Clelia errabunda sp. nov., BMNH 89.8.14.12. Dissection to 
show superficial head muscles and glands (reversed). 
Cc = constrictor colli muscle, overlaying other structures; 
Cm = cervico-mandibular muscle; Dg = Duvernoy's gland; 
Hg = Harder's gland; Ig = infralabial gland; 
Ql = quadrato-labial ligament; Sg = supralabial gland; 
Rg = rictal gland. 

brazili and unlike atrox. In all, the dorsal bands are indistinct. 
In two of the Demerara specimens the bands can be seen to 
have parallel sides or to converge towards the dorsal midline 
as described by Lazell for caribbaeus. The third shows some 
diverging bands. The Guyana atrox have dark patches on the 
lower flanks which extend onto the ventrals unlike these 
Sabine specimens. The brazili have dark bordered bands 
which converge towards the midline. These observations 
suggest that the 'Berbice' and 'Demerara' specimens are most 
probably caribbaeus, a species known only from St Lucia. 
The above considerations raise a doubt that Sabine collected 
any specimens in Guyana. 

It is evident that Sabine was long enough, supposedly in 
Guyana, to achieve promotion from captain to colonel; in 
that time he may well have visited St Lucia. The locality of 
the 'Demerara' Clelia errabunda is therefore discounted. The 
uniform 17 scale-rows and undifferentiated vertebrals would 
distinguish it from local Clelia clelia. 

Dumeril, Bibron & Dumeril (1854) report that the Paris 
museum has specimens of 'Brachyruton cloelia from Guy- 
ana, Brazil, Mexico and Guadeloupe. The Guadeloupe' 
specimen (MNHP 169) is a hatchling with counts V200, C83 
and 19:19:19 scale rows. The tip of the heart is at V46, the 
trachea extends beyond V90. This is clearly a specimen of 
Clelia clelia, with an unusually low ventral count. The locality 
is undoubtedly erroneous. 

In the Proceedings of the Philadelphia Academy for 1870 it 
is reported that Cope "called attention to a large specimen of 
Trigonocephalus ( = Bothrops) from St Lucia, of which some 
fourteen inches was enclosed in the oesophagus and stomach 
of a larger Oxyrhopus plumbeus ( = Clelia clelia)." Later 
Cope (1876) wrote that, as he had previously observed, he 
had received a specimen of Clelia clelia from Martinique(l) 
which had swallowed a large Bothrops. Malnate (personal 
communication) examined the specimen(s). It is ANSP 10220 
from 'Santa Cruz', received from Mrs J.L. Endicott. 'Santa 
Cruz' presumably means St Croix in the Virgin Islands! It is 
unlikely to be a St Lucia locality; most of the place names are 
French. This one specimen thus has three different geograph- 
ical attributions! 

Malnate reports that the Clelia has 17 scale-rows through- 
out and an undifferentiated vertebral scale row; it therefore 
fits C. errabunda. The half-swallowed Bothrops has 25 scale 
rows about midbody, falling to 19 rows. Lazell (1964) gives 



G. UNDERWOOD 



mid-body scale-row counts for Bothrops caribbaeus from St 
Lucia as 25-29 (mode 27) and for Bothrops lanceolatus from 
Martinique counts from 29-33 (mode 31). The Philadelphia 
Academy specimen, with 25 rows, is at the lower end of the 
range for St Lucia specimens. Beyond reasonable doubt 
therefore Cope had a specimen of C. errabunda which had 
half swallowed a St Lucia Bothrops caribbaeus. 

Relationships of Clelia errabunda 

The species currently assigned to Clelia can be arranged at 
several levels on the basis of the derived states of the 
respiratory system, the vertebral scale row and the ventral 
scale counts. This is set out in Table 3. 

C. bicolor. The three anterior supralabials, the low ven- 
tral scale counts, the undifferentiated vertebral scales, the 
presence of a left lung and a short trachea are primitive 
features. The high maxillary tooth count appears to be a 
derived feature. 

C. rustica. This species also has undifferentiated verte- 
bral scales, a left lung (a mere vestige in BMNH 81.7.2.9) and 
a short trachea. C. rustica and the following species are 
derived in relation to bicolor in respect of two anterior 
supralabials and higher ventral counts. 

C. errabunda, clelia, equatoriana and scytalina share the 
derived feature of absent left lung. The latter three species 
are further derived than errabunda in respect of the enlarged 
vertebral scales and extended trachea. 

The type specimen of Clelia clelia Daudin (1803) is not 
known to survive; however, the type locality is given as 
'Suriname'. It is therefore assumed that specimens from the 
northern coast of South America are typical clelia. Specimens 
in The Natural History Museum from this area, from Central 
America, from Rio Condoto on the Pacific slope of Colombia 



(1), from La Paloma nr Santiago R., Ecuador (1) and from 
most of the rest of South America show the 19:19:17 scale 
row pattern. However, a second specimen from Rio Condoto 
and specimens from Ecuador (Guayaquil and east of Loja, 
2), Peru (3) and Manacapuro on the Amazon (1) show the 
17:19:17 scale-row pattern. These are indistinguishable from 
typical clelia in respect of the other characters considered 
here. Their occurrence in a north-western area of South 
America with near overlap with the 19:19:17 (Rio Condoto) 
form on the Pacific slope of Colombia does not look like an 
accident of sampling (Fig. 5). The form in eastern Brazil 
(plumbed) lacks spines on the hemipenis, some specimens 
from the southern part of the range (occipitolutea) are pale in 
colour, so clelia is evidently a variable species. Roze (1959) 
reports a specimen from Venezuela with counts of: 
21:22:19:17 and Chippaux (1986) reports a specimen with 
21:19:17 rows from French Guyana. 

After the above account was prepared I received from 
Zaher (personal communication) a photocopy of a portion of 
Bailey's unpublished PhD thesis. It is evident that at that time 
he regarded the island Clelia as sufficiently distinct to merit 
subspecific status. He too did not believe the Guyana locali- 
ties of the Sabine specimens. 



DISCUSSION 

The species clelia, equatoriana, scytalina and errabunda share 
uniform dark adult colouration and absence of a small left 
lung. Most other pseudoboine snakes have a small left lung 
and a more varied colour pattern. Within this group of four 
species errabunda is primitive to the others in that the 
vertebral scale row is not enlarged and the trachea is short. 




Fig. 5. Localities of specimens from northern South America and the Lesser Antilles. Solid symbols = precise localities; hollow 
symbols = approximate localities; circles = Clelia clelia; triangle = Clelia errabunda; 17, 19 = no of scale rows on neck of C. clelia. 
B = Barbados, D = Dominica, Gr = Grenada, Gu = Guadeloupe, L = St Lucia, M = Martinique, V = St Vincent. 



A NEW SNAKE FROM ST LUCIA 



Because the 19:19:17 scale row pattern is widespread in 
pseudoboine snakes and is also found in most clelia we may 
infer that it was the condition of the ancestor of this species 
group. The 17:17:17 pattern shown by errabunda would thus 
be interpreted as a derived feature setting it apart. However, 
the occurrence of the 17:17:17 pattern in equatoriana and 
scytalina and the 17:19:17 and 21:19:17 patterns within clelia 
suggests that little significance can be attached to the scale- 
row pattern. The most nearly special feature of the new 
species is the high number of large spines on the hemipenis. 
Otherwise it is close to the status of what Ackery and 
Vane-Wright (1984) call a 'paraspecies', without any special 
feature setting it apart. The short trachea and unmodified 
vertebral scales by which it is distinguished are primitive 
features found in hundreds of other species of snakes. 

Greer (1965) reports that the Grenada Clelia is diurnal, 
unlike its mainland relatives. Clelia clelia from Grenada is 
otherwise little different from mainland clelia; this is con- 
firmed by Wallach's report (personal communication) that it 
has an extended trachea. It is presumably a relatively recent 
immigrant from South America. On the other hand erra- 
bunda, on St Lucia, is primitive to the mainland members of 
the clelia group. This suggests that it colonised St Lucia at an 
early date and that its ancestral stock was later replaced on 
the mainland by the more derived clelia. It is an example of a 
primitive form surviving as an island relict. 

Boa constrictor occurs on St Lucia and Dominica. The two 
island populations and the mainland form are well differenti- 
ated from one another and are recognised as separate subspe- 
cies (Lazell, 1964). The pit-vipers, Bothrops, on the adjacent 
islands of St Lucia and Martinique are sufficiently differenti- 
ated that they are recognised as full species by Lazell (1964). 
For both Boa and Bothrops this suggests either separate 
colonisation of the islands from the mainland or colonisation 
of one island and passage to the other long ago. There is at 
present no evidence that these are primitive island relicts. 

Cope (1870) is reported as saying that the "islands of 
Martinique and Guadeloupe had become so infested with the 
fer-de-lance" {Bothrops lanceolatus) "as to be in parts almost 
uninhabitable, and it was chiefly on account of the danger 
from this venomous reptile that collecting naturalists of late 
years had so seldom visited them"! "Some means had been 
adopted to check the increase of this pest, but with small 
results". "Prof Cope thought that as the Oxyrhopus 
plutnbeus (= Clelia clelia) was very numerous in Venezuela 
and Brazil, and since it was very harmless and easily pro- 
cured, that its introduction in large numbers into Martinique, 
etc, would be a simple matter, and one probably to be 
attended with good results in the diminution, at least, of this 
enemy of agriculture". 

Lazell (1964) tells us that on both Martinique and St Lucia 
the local Bothrops is known as 'serpent'. We may speculate 
that prior to human arrival 'serpents' were already estab- 
lished on St Lucia before the 'cribo' (Clelia) arrived to prey 
upon them. 

It is said that the mongoose was introduced into St Lucia in 
the hope that it would reduce the Bothrops. Today, although 
the mongoose may eat Bothrops, it also eats domestic poul- 
try. Following human disturbance, it is ironic that the indige- 
nous 'pest', the 'serpent', is supplemented by an introduced 
pest, the mongoose, and the indigenous biological control, 
the 'cribo', is extinct! In the absence of this 'control' Lazell 
(1964) reports that in some areas of St Lucia the serpent is 
'abundant beyond belief. 



Acknowledgements. I am indebted to Van Wallach for a report on 
a Grenada specimen of Clelia clelia and for information about other 
species of Clelia and other xenodontine snakes. Beat Schatti loaned a 
specimen of Clelia equatoriana for examination of the anterior 
viscera. Ed Malnate reported on the Philadelphia Academy speci- 
men^). Robert Henderson, David Bullock and Roger Thorpe 
replied to a query about the status of Clelia on Dominica. David 
Corke commented on the status of Clelia on St Lucia. Hussan Zaher 
drew my attention to the work of Scrocchi and Vinas, identified the 
Oxyrhopus trigeminus from 'Guyana', located specimens in the Paris 
Museum and gave me a photocopy of a portion of Bailey's unpub- 
lished PhD thesis. T.E. Pickring, archivist at The Natural History 
Museum, traced the early records of Capt Sabine. Colin McCarthy 
helped me to find my way through The Natural History Museum 
records and read a first draft of this paper. The Museum national 
d'Histoire naturelle, Paris, loaned two specimens. 



REFERENCES 



Ackery, P.R. & Vane-Wright, R.I. 1984. Milkweed butterflies, their Cladistics 
and Biology. British Museum (Natural History), London. 

Barbour, T. 1930. A list of Antillean reptiles and amphibians. Zoologica, 11: 
61-116. 

1935. Second list of Antillean reptiles and amphibians. Zoologica. 19: 

77-141. 

1937. Third list of Antillean reptiles and amphibians. Bulletin of the 

Museum of comparative Zoology 82: 1-166. 

Bullock, D.J. & Evans, P.G.H. 1988. The distribution, density and biomass of 

terrestrial reptiles in Dominica, West Indies. Journal of Zoology. 222: 

421-443. 
Boulenger, G.A. 1896. Catalogue of the Snakes in the British Museum, Vol. 3, 

part A. 
Cadle, J.E. 1984. Molecular systematics of Neotropical Xenodontine snakes: I 

South American Xenodontines. Herpetologica. 40: 8-20. 
Chippaux, J-P. 1986. Les serpents de la Guyane franchise. Editions de 

l'Orstom, Institut fran^ais de recherche scientifique pour le developpment en 

cooperation. Collection fauna tropicale. no XXVII, Paris. 
Cope, E.D. 1870. Verbal communication' Aug 2nd. In: Proceedings of the 

Academy of Natural Sciences of Philadelphia, p. 90. 

1876. On the Batrachia and Reptilia of Costa Rica. Journal of the 

Academy of Natural Sciences. Philadelphia, 8(2), p. 131. 

Dowling, H.G. 1951. A proposed standard system of counting ventrals in 

snakes. British Journal of Herpetology, 1: 97-99. 
Dumeril, A.-M.-C, Bibron, G. & Dumeril, A. 1854. Erpetologie generate. 

Paris. Vol. VII, part II, p. 1007. 
Greer, A. 1965. A new subspecies of Clelia clelia (Serpentcs, Colubridae) from 

the island of Grenada. Breviora, 223: 1-6. 
Haas, G. 1973. Muscles of the jaws and associated structures in the Rhyn- 

chocephalia and Squamata. In: Gans, C. & Parsons, T.S. (eds) Biology of the 

Reptiles. Academic Press, London. 
Lazell, J.D. 1964. The Lesser Antillean representatives of Bothrops and 

Constrictor. Bulletin of the Museum of comparative Zoology, 132 (3): 

245-273. 
Long, E. 1974. The serpent's tale. U.W.I. Extra mural department, P.O. Box 

306, The Morne, St Lucia. 
McDowell, S.B. 1986. The architecture of the corner of the mouth of colubroid 

snakes. Journal of Herpetology, 20: 353-407. 
Peters, J.A. & Orejas-Miranda, B. 1970. Catalogue of the Neotropical Squa- 
mata: Part I, Snakes. United States national Museum Bulletin, 297, Washing- 
ton. 
Roze, J.A. 1959. Taxonomic notes on a collection of Venezuelan reptiles in the 

American Museum of Natural History. American Museum Novitates, No 

1934: 1-14. 
Schwartz, A. & Henderson, R.W. 1988. West Indian amphibians and reptiles: a 

checklist. Milwaukee Public Museum, Bulletin 74, Wisconsin. 
Scrocchi, G. & Vinas, M. 1990. El genero Clelia (Serpentes: Colubridae) en la 

Republica Argentina: revision y comentarios. Bolletino del Museo regional 

dei Scienze naturale di Torino, 8: 487-499. 
Smith, M.A. 1943. Fauna of British India. Reptilia and Amphibia, Vol. 3, 

Serpentes, Taylor & Francis, London. 
Smith, M.A. & Bellairs, A.d'A. 1947. The head glands of snakes, with remarks 

on the evolution of the parotid gland and teeth of the Opisthoglypha. Journal 



8 



G. UNDERWOOD 



of the Linnaean Society, Zoology, 61: 351-368. 
Thorpe, R.S. 1975. Quantitative handling of characters useful in snake 

systematics with particular reference to intraspecific variation in the ringed 

snake Natrix natrix (L.) Biological Journal of the Linnaean Society, 7: 27-43. 
Underwood, G. 1963. A contribution to the classification of snakes. British 

Museum (Natural History), London. 



Underwood, G. & Stimson, A.F. 1990. A classification of pythons. Journal of 
Zoology, 221: 565-603. 

Vanzolini, P.E. 1986. Addenda and corrigenda to the catalogue of neotropical 
Squamata. Smithsonian herpetological information service, No 70, Washing- 
ton. 



APPENDIX 



Table 1 Some representative specimens of Clelia, showing: sex, ventrals, scale row reduction pattern, scale row stretches on tail, 
specimen of Oxyrhopus maculatus Boulenger. 



= type 



Tail row stretches 







V 


Row reductions 


scytalina 

68.4.7.7 


M 


212 


»19(12;5>)17 


68.4.7.8 


F 


209 


>»19(7;5>)17 


equatoriana 








60.6.16.47 


M 


201 


»17 


Geneva 2410.9 


M 


201 


»17 


60.6.16.48 


F 


217 


»17 


60.6016.49 


F 


219 


»17 


clelia 








74.8.4.56 


M 


216 


»17(44;5<)19(149;5>)17 


90.10.6.29 


M 


214 


>»17(41;4<)19(146;5>)17 


1926.4.30.14 


F 


238 


»17(43;4<)19(159;4>)17 


51.7.17.136 


M 


219 


>19(165;4>)17 


86.10.4.12 


M 


213 


»19(165;4>)17 


1929.10.19.2 


F 


237 


>19(209;5>)17 


1902.7.29.68 


F 


231 


»19(142;6>)17 


* 84.2.23.40 


F 


213 


»19(179;5>)17 


errabunda 








89.8.14.25 


M 


221 


»17 


MHNP 7598 


M 


224 


»17 


89.8.14.12 


F 


234 


»17 


1988.717 


F 


230.5 


»(7;4>)17 


rustica 








86.1.19.21 


M 


206 


»19(176;5>)17 


81.7.2.9 


M 


196 


>»19(129;4>)17 


95.9.17.21 


F 


212 


»19(177;4>)17 


1933.9.5.7 


F 


195 


»19(128;4>)17 


bicolor 








1927.8.1.234 


F 


178 


19(96;5>)17 


1980.1651 


F 


177 


19(108;4>)17 



>8 



75 


15 


11 


20 


25 


4 


83 


11 


14 


22 


30 


6 


75 


5 


17 


23 


30 





69 


4 


14 


18 


28 


5 


- 


3 


12 


20 


- 


- 


- 


5 


12 


21 


- 


- 


91 


10 


19 


26 


26 


10 


71 


6 


20 


27 


18 





82 


2 


8 


24 


26 


12 


91 


6 


20 


24 


33 


8 


64 


12 


13 


19 


17 


3 


84 


3 


14 


32 


30 


5 


55 


3 


7 


25 


20 





50 


3 


11 


20 


19 





84 


11 


15 


22 


34 





75+ 


15 


14 


21 


25+ 


- 


71 


5 


8 


27 


31 





- 


4 


11 


- 


- 


- 


61 


14 


22 


16 


9 





60 


6 


23 


17 


16 





55 


7 


17 


19 


12 





39 


3 


9 


21 


6 





59 


3 


15 


15 


21 


5 


58 


3 


12 


16 


23 


4 



A NEW SNAKE FROM ST LUCIA 

Table 2 Distribution of tubercles on the dorsal, frontal and parietal scales of some selected specimens of Clelia. 



Dorsal scale-row nos. 



10 



par 



scytalina 


68.4.7.7 


equatoriana 


60.6.16.47 


clelia 19:19:17 


1930.10.10.188 


clelia 19:19:17 


94.3.14.60 


clelia 17:19:17 


89.4.8.2 


errabunda St L. 


89.8.14.25 


errabunda St L. 


MNHP 7598 


errabunda 'Dominica' 


89.8.14.12 


errabunda 'Demerara' 


1988.719 


rustica 


1909.11.2.16 


bicolor 


1980.1651 



M--4142154 
M - 1 - - 1 1 1 1 2 
M-------23 2 

F 1 3 1 14 2 4 4 3 4 
M23- 233346 6 
M-----12-- 

M-----12-- 

F----34532 

F---11112- 
M---------- 

F ----- 1 1 1 1 1 



117 


98 


129 


98 


151 


104+ 


96 


87 


150 


112 


138 


92 


200 


104 


186 


152 


141 


132 


72 


45 + 


45 


34 



Table 3 Comparison of Clelia species. 















Max 






Vert. 






n 


suplabs 


V 


C 


teeth 


L.lung 


Trachea 


row 


scytalina 


F 


1 


2:2:3 


209 


83 


13 


+ 


+ 


+ 




M 


1 




212 


75 


14 








equatoriana 


F 


2 


2:2:3 


217-219 


- 


11.5 


+ 


+ 


+ 




M 


2 




201 


72 


12 








clelia 


F 


15 


2:2:3 


213-228-238 


50-73-84 


11.5 


+ 


+ 


+ 




M 


13 




204-215-226 


64-82-91 


11.3 








errabunda 


F 


2 


2:2:3 


231-234 


71 


14 


+ 


- 


- 




M 


2 




221-224 


84 


13.7 








rustica 


F 


7 


2:2:3 


195-208-231 


39-44-55 


11.3 


- 


- 


- 




M 


2 




195-210-223 


61 


13.0 








bicolor 


F 


2 


3:2:3 


177-178 


58-59 


15.5 


- 


- 


- 



V = ventrals, C = subcaudals, min-mean-max; mean no of anterior maxillary teeth; - = primitive, + = derived state of left lung, trachea and vertebral scale-row. 



Bull. nat. Hist. Mus. (Zool.) 59(1): 11-31 



Issued 24 June 1993 



Anatomy of the Melanonidae (Teleostei: 
Gadiformes), with comments on its 
phylogenetic relationships 

GORDON J. HOWES 

Department of Zoology, The Natural History Museum, Cromwell Rd, London SW7 5BD 

CONTENTS 



Introduction 11 

Materials and methods 12 

Abbreviations used in the figures 12 

Anatomy 13 

Neuromast pattern 13 

Infraorbitals 14 

Cranium 16 

Jaws 19 

Palatopterygoquadrate 20 

Hyoid arch 21 

Opercular bones 22 

Branchial arches 22 

Pectoral girdle 23 

Pel vie girdle 24 

Vertebral column and median fins 25 

Brain 27 

Swimbladder, viscera and body musculature 28 

Discussion 28 

Acknowledgments 30 

References 30 



Synopsis. The osteology and part of the soft anatomy of the gadiform family Melanonidae, represented by the 
genus Melanonus Giinther, 1878, is described. Melanonus has several derived (autapomorphic) sensory features but 
only three osteological ones. Although contained within the Gadiformes the family is excluded from both the 
Macrouroidei and Gadoidei in lacking a modified palatine and enlarged intercalar and thus represents their 
sister-group designated as the Melanonoidei. The Suborder Gadoidei now comprises two families of uncertain 
phylogenetic affinity (Bathygadidae and Steindachneriidae) and two Superfamilies, Moridoidea and Gadoidea. 



INTRODUCTION 



The gadoid genus Melanonus Giinther 1878 contains two 
species, M. zugmayeri Giinther, 1878 (Fig. 1), and M. gracilis 
Norman, 1930, which together give a broad latitudinal, 
circumglobal distribution (Cohen et al., 1990; Howes, 1991a). 
Melanonus are relatively small fishes, the largest seen being 
230mm total length and, oddly for gadoids, are meso- 
bathypelagic (100-3000m). Outwardly, Melanonus resembles 
a stomiatoid rather than a gadoid fish with its dark coloration, 
large, strongly-toothed jaws and tapering body (Fig. 1). 

Until Marshall (1965) recognised (without diagnosis) a 
separate family for the genus, Melanonus had been consid- 
ered to belong to the Moridae. Marshall (1965) and Marshall 
& Cohen (1973) contended that Melanonus was the most 
primitive gadiform (anacanthine) fish, a contention based on 



the posterior position (at the forebrain) of the olfactory bulbs 
and a relatively unmodified caudal fin skeleton. The features 
Marshall & Cohen (1973) used to diagnose the Melanonidae 
rested on soft anatomical features, viz. an elaborate system of 
free ending neuromasts on the head and the corpus cerebelli 
extending (forward) to the optic tectum. 

Apart from a few observations on the caudal fin skeleton 
and gill-arches and a description of its cranial muscles (see 
below) the anatomy of Melanonus has never properly been 
described. Gosline (1971) complained that "No account of 
the osteology is available. By contrast the family Gadidae has 
received more attention from anatomists than almost any 
other family of fishes". Despite these shortcomings several 
assertions as to the phylogenetic position of the Melanonidae 
have been made. 

Rosen & Patterson (1969) cited Marshall (1965; 1966) to 
the effect that Melanonus represents a primitive gadoid. 



12 



G.J. HOWES 



















JR &i 






















•■•'.' '■■■ 


^ 




























II J * 










5*g . 


^B 


^? 




^fl 


■ 


^^^^5 




* 




^Hb \ 


^aJ 


wm. 


















■•/ ■ 




^^gMM 






^L v 


- B 










■V.'/ 












_^^^b 


















fig%£ 




**J 


hi! 








--iaitt*^ 












































r^^^K 






H^^H^^I 








IPJUli 


pupil 


II 


r 






■K^^HrjlHt 


i-HE^lB^^ 





Fig. 1. Melanonus zugmayeri specimen BMNH 1991.7.9:729-30, 200mm SL, lateral view. 



Schwarzhans (1980; 1984) combined the Melanonidae and 
Moridae (produced as a cladogram in Patterson & Rosen, 
1989) and Cohen (1984) and Fahay & Markle (1984) also 
suggested a relationship with the Moridae, again based on the 
primitive arrangement of the caudal fin skeleton. Markle 
(1989) revised his earlier views and placed the Melanonidae 
near the base of his cladogram making it (with the exception 
of the Ranicipitidae) the sister group of all other gadiforms. 
Nolf & Steurbaut (1989) placed Melanonidae as an unre- 
solved polychotomy with the Euclichthyidae, Macrouridae, 
Moridae and other gadoids. Okamura (1989) omitted the 
family from his gadoid classification but implied (p. 137) on 
the basis of similar anterior rib structure that Melanonus is 
closely related to Merluccius. Howes (1989; 1990) also placed 
Melanonidae in an unresolved polychotymy, with Stein- 
dachneriidae, Bathygadidae and other gadoids. According to 
Howes (1990, 1991a & b) the majority of gadoid families 
form a monophyletic assemblage termed 'supragadoids', 
characterized by complete fusion of the upper hypurals into a 
single plate. The Macruronidae represent the plesiomorphic 
lineage of this assemblage with the Gadidae and Merlucciidae 
being the most derived families. The Melanonidae was 
assigned with the Bathygadidae, Steindachneriidae, Moridae 
and Euclichthyidae to the 'infragadoids' and in one scheme 
(Howes, 1991b) in alternative positions, one as the sister 
group to all gadoids excluding the Bathygadidae and Stein- 
dachneriidae, the other as also excluding the Moridae. The 
characters on which these phylogenetic positions were based 
are, however, ambiguous (see Discussion) and like all previ- 
ous studies have suffered from lack of anatomical information 
about Melanonus. The following is an account of the osteol- 
ogy and other soft anatomical features of Melanonus. 



MATERIALS AND METHODS 

Specimens used for anatomical descriptions (all from BMNH 
collections): Melanonus zugmayeri Uncat. 230mm, 'Discov- 
ery' Stn 11550, 10.VII.1987, 20°25.5'N, 19°39.5'W, 
775-825m; 1991.7.9:729-730, 220mm, 95mm SL, 
20°25.8'N-31.4'N, 19° 39.8'W-38.0'W, 800-875m; 
1991.7.9:731-733, cleared & stained, 66, 100, 130mm SL, 
17°1.2'N, 19°57.8'W, 400-495m; 1987.1.21:595-596, 215mm 



SL, dry skull prepared from 190mm SL, 49°21.9'N, 11°51'W, 
1090-llOOm; 1987.1.21:597, 168mm SL, S.W. Bantry, 
960-920m; 1981.3.16:377, 173mm SL, West Great Sole 
Banks; 1987.1.21:598-601, 175, 187, 193mm SL, one speci- 
men, 165mm SL (cleared and stained), 50°02'N, 11°22'W, 
910-980m; 1930.1.12:943 (Holotype) 13°58'S, 11°43'E. Mel- 
anonus gracilis 1887.12.7:22 (Holotype) 147mm SL, Antarc- 
tic; 1930.1.12:934-936, 97, 140, 150mm SL, 46°56'S, 
46°03'W; 1988.11.4:13-20, 45, 49mm SL (cleared and 
stained), 35°13'-34°57'S, 17°50'-17° 48'E; 1988.11.4:2, 
145mm SL, 50°17.7'S, 18°40.9'E, 300-150m; Percopsis omis- 
comayus 1973.3.20:46-8, 52mm (cleared and stained), 62mm 
SL, Lac Henry, Quebec, Canada; Bregmaceros sp. 
1957.12.2:5-12, 54mm SL (cleared and stained) Senegal; 
Gaidropsarus mediterraneus Uncat. 122, 145mm (cleared and 
stained), Seaton Point, England. 

In addition, material listed in Howes (1988, 1992) and 
Howes & Crimmen (1990) was re-examined. 

Abbreviations used in the figures 



aa 


anguloarticular 


aap 


premaxillary articular process 


ac 


actinost 


afc 


anterior frontal crest 


ap 


premaxillary ascending process 


ar 


anterior ('chopstick-like') ribs 


ard 


anal fin radial 


asp 


autosphenotic 


bb 


basibranchial (numbered) 


bh 


basihyal 


bl 


Baudelot's ligament 


bo 


basioccipital 


boc 


basioccipital condyle 


br 


branchiostegal ray 


cb 


ceratobranchial (numbered) 


cc 


cerebellar corpus 


ccr 


cerebellar crest 


cfc 


central frontal crest 


cl 


cleithrum 


cm 


coronomeckelian bone 


CO 


coracoid 


de 


dentary 


dex 


dorsal section of epaxialis muscle 


dhy 


dorsohyal 


dr 


dorsal fin ray 



ANATOMY OF THE MELANONIDAE 



13 



dorsal fin radial 

epibranchial (numbered) 

epibranchial toothplate 

ectopterygoid 

erector and depressor dorsalis muscles 

entopterygoid 

epural 

epineural 

extrascapular 

epioccipital 

exoccipital condyle 

exoccipital cartilage 

exoccipital flange 

exoccipital 

foramen for glossopharyngeal nerve 

foramen for vagus nerve 

facial lobe 

foramen magnum 

foramen for occipital nerves 

frontal 

flexor vert trails muscle 

flexor ventralis inferioris muscle 

granular eminence 

gas-gland 

gonad 

gut 

haemal arch 

hypobranchial (numbered) 

hyomandibular fossa 

hypophysis 

hypural (numbered) 

interarcual cartilage 

intercalar 

interhyal 

infraorbitals (numbered) 

interopercle 

interradialis muscle 

kidney 

lateral ethmoid 

liver 

lateral ethmoid-palatine ligament 

mandibular-interopercular ligament 

trigeminal lobe 

Meckel's cartilage 

mesonephric duct 

mesethmoid cartilage 

metapterygoid 

myoseptal strands supporting ribs 

medulla oblongata 

cranial nerves 

nasal 

first neural arch 

nerve branch serving adductor arcus palatini muscle 

acoustic nerve 

buccalis branch of trigeminal trunk 

infraorbital branch of trigeminal trunk 

lateral line nerve 

neuromasts types 1 and 2 

nerve of supraorbital branch innervating posterior canal 

enclosed neuromast 

nasal rosette 

supraorbital branch of trigeminal 

olfactory bulb 

opercle 

olfactory tract 

parietal 

parhypural 

palatine 

pelvic bone 

pharyngobranchial (numbered) 



pc 


postcleithrum 


pfc 


posterior (diagonal) frontal crest 


phy 


posterohyal 


pmp 


postmaxillary process of premaxilla 


pop 


preopercle 


PP 


parapophysis 


prn 


prootic notch 


ps 


parasphenoid 


psl 


parasphenoid ascending laminae 


pte 


pterotic 


pts 


pterosphenoid 


ptt 


posttemporal 


pu 


preural vertebra (numbered) 


pyc 


pyloric caeca 


qu 


quadrate 


ra 


retroarticular 


Rcl 


ramus canalis lateralis nerve 


rd 


retractor dorsalis muscle 


re 


rostrodermosupraethmoid 


sb 


swimbladder 


sbp 


swimbladder pocket 


sc 


scapular 


sea 


supracarinalis anterior muscle 


scl 


supracleithrum 


so 


supraoccipital 


sop 


subopercle 


spt 


intercalar socket for posttemporal limb 


St 


stomach 


sy 


symplectic 


ul 


ural centrum 


V 


vertebra (numbered) 


vex 


ventral section of epaxialis muscle 


vhy 


ventrohyal 


vo 


vomer 



ANATOMY 



Neuromast pattern (Fig. 2). 

Melanonus has a unique pattern of free-ending neuromasts 
covering the head in addition to those more usual neuromasts 
contained in the sensory canals. There are two morphotypes 
of the former: 1) the most numerous, are long, flange-like 
structures which occur on the skin covering the upper rim of 
the infraorbitals, the snout, cheek muscles, preoperculum 
and top of the head; 2) button-like structures confined to 
specific areas on the lower cheek, snout and frontal. 

The flange-like neuromasts are usually arranged longitudi- 
nally and more or less in rows. In the snout region, individual 
neuromasts may be slightly curved or angled to the general 
direction of the others (Fig. 2B). On top of the head, rows are 
more definite and those on the snout tend to converge 
anteriorly where the organs close to the midline are larger 
than the others (exceptional is the star-shaped arrangement 
posterior to the medial extrascapular sensory pore in M. 
zugmayeri); the neuromasts close to the midline on the 
central part of the frontal are also nearly twice the length of 
the others (Fig. 2A). In M. zugmayeri the neuromasts along 
the anterior part of the supraoccipital have a regular arrange- 
ment (Fig.2A) but in M. gracilis they form a pocket or 
enclosed area. Distribution on the preoperculum is irregular 
and sometimes sparse (the skin is often missing from this 
region and it is not possible to make precise counts). 

The pitlines of button-like organs are in a double row near 
the border of the snout, in a patch above the nares, as an 



14 



G.J. HOWES 



nml 




' 'V'/.V iflll 

i. /' , n ; \i' 

M ''Hi • 1 1 -| M/ 

// ///// I 1 ,' # 
I I J ' I 1 , •» /' ' 

' *-. / ' ' 'i/l / 

i \ w / 











-\ \ 






\ 



X 



-3 ^ 



l 

Fig. 2. Distribution of neuromasts types 1 and 2 in A, Melanonus zugmayeri on dorsal surface of head and B, M. gracilis on lateral surface 
of head. C, innervation pattern of type 1 neuromasts in subnasal region (right side) of M. zugmayeri (dashed lines indicate nerves, arrow 
heads indicate termination of nerve branch; large arrow points anteriorly). In this and subsequent figures, scale bars in millimetre divisions. 






oblique row across the lower part of the cheek and a double 
row across the epioccipital region (Figs 2A,B). The neuro- 
masts do not extend backwards on to the body. 

There are about 500 flange-like neuromasts covering the 
entire head. Innervation is by the ramus canalis lateralis (Rcl 
sensu Freihofer, 1970) which exits from the posterior frontal 
foramen to anastomose through a loose fascia of connective 
tissue. The neuromasts are innervated by subranches stem- 
ming from a complex nerve network (Fig. 2C). 

The Rcl nerve branches from the supraorbital trunk of the 
trigeminal complex, (Fig. 17), a condition similar to that in 
Merluccius (Freihofer, 1970). The branch innervating the 
large posterior neuromasts enclosed in the frontal sensory 
canal detaches separately from the supraorbital trunk, 
whereas in Merluccius the nerves separate off together. 

The large, plate-like neuromasts, housed in the sensory 
canals number two in the nasal bone, three in the frontal (one 
beneath the anterior medial ridge, one beneath the lateral 
arch and one posteriorly), one in the anterior part of the 



pterotic, one in the parietal, one in each extrascapular, three 
in the first infraorbital, one in the second, third and fourth, 
two in the fifth and one in the sixth, and five in the 
preoperculum. 

Infraorbitals (Fig. 3). 

There are six infraorbital bones, the first long and relatively 
deep with a broadly fretted ventral border, the outer flange 
which forms the roof to the sensory canal extends as a shelf 
along the anterior half of the bone, but curves laterally along 
the posterior half. The ascending process which contacts the 
posterior wall of the lateral ethmoid is tall and spine-like. The 
second infraorbital is confluent with the first and is as deep 
but only a sixth of its length. The medial lamina of the third is 
widely separated from that of the second although the 
dorsolateral flange is nearly in contact. The third together 
with the fourth form the posteroventral corner of the orbit 
and the fourth has only a short orbital margin and flange 



ANATOMY OF THE MELANONIDAE 



15 




Fig. 3. Infraorbital bones of M. zugmayeri in specimens of: A, 66mm SL; B, 100mm SL and C, 130mm SL. 



posteriorly, the body of the bone is expanded. The fifth 
infraorbital has a long orbital margin, the lower part of which 
projects anteroventrally in front of both the third and fourth 
to which it is connected by strong connective tissue; it has a 
narrow flange along its upper orbital border. The sixth 
(dermosphenotic) is as large as the fifth and has a pronounced 
orbital curvature which brings its anterior tip to the same 
vertical plane as the ascending process of the first infraor- 
bital. 

In the two smaller specimens of M. zugmayeri examined, 
the dorsolateral flange remains undeveloped on the first, fifth 
and sixth infraorbitals of the 66mm specimens and the 
ascending process of the first infraorbital is inclined anteriorly 
in both (Fig. 3A). The anteroventral border of the fifth 
infraorbital is less pronounced and in the 100mm SL speci- 
men its tip lies medial to the rim of the fourth infraorbital; the 
sixth lacks the anterior elongation of the larger (130mm SL) 
specimen (Figs 3B, C). 



Unlike other gadiforms where the posterior (fifth and 
sixth) infraorbitals are shallow, those of Melanonus are as 
deep as the anterior ones. The anterior curvature of the 
upper infraorbital (dermosphenotic) is more reminiscent of 
some macrouroids (see below) than gadoids. The central 
position of the ascending process of the first infraorbital is 
probably a plesiomorphic gadiform feature (on the basis of 
commonality) as is the reduced size of the second infraor- 
bital. The extension of the lower part of the fifth infraorbital 
and the enlargement of the dermosphenotic are, because of 
their restricted distributions, taken to be derived features. 
According to Iwamoto (1989) among macrouroids, exclusion 
of the third and fourth infraorbitals from the orbit is a derived 
condition. In Melanonus similar exclusion has resulted from 
ventral extension of the fifth infraorbital whereas in the 
macrouroids illustrated by Iwamoto (1989, fig. 5G) it is due 
to re-alignment of the fourth infraorbital which covers the 
orbital borders of the second and third. 



16 



G.J. HOWES 



me 




esc 



so 



Fig. 4. Neurocranium of M. zugmayeri in A, dorsal and B, ventral views. In A, the right parietal, right nasal and left posttemporal have 
been removed. In B, dashed outline circles on the prootic and intercalar indicate the positions of the otoliths. 



Cranium (Figs 4-8). 

In its overall shape the cranial roof is almost square, the most 
noticeable feature being the deep indentation of the lateral 
frontal border anterior to the sphenotic, and the prominent 
anterolateral projections of the lateral ethmoid wings 
(Fig. 4A). 

The ethmoid dorsal surface (rostrodermosupraethmoid) is, 
in keeping with that of other gadoids (Howes & Crimmen, 
1990: 166), being narrow and cruciform with a steep anterior 
slope (Fig. 5B). The ossified anterior wall of the ethmoid 
forms most of the nasal cavity and a thin, vertical septum of 
ethmoid cartilage separates the cavities medially. A shallow 
bed of cartilage separates the base of the ethmoid and the 
vomer. The vomer has a thick, broadly rounded head bearing 
on either side 6 or 7 teeth in smaller specimens and 10-12 in 
larger (Figs 4B, 6A). In smaller specimens of both species the 
teeth are more or less arranged in a single row but in larger 
specimens the posterior teeth tend to be in a patch with one 
or two stout and caniniform being almost twice the length of 
their neighbouring teeth and three times that of the symphy- 
seal teeth (Fig. 6A). The vomerine shaft is relatively short, 
extending to just beyond the posterior level of the lateral 
ethmoid. The base of the lateral ethmoid is long and broad 
and where it meets the vomer bears a deep cavity into which 
inserts the palatine ligament. The wall of the lateral ethmoid 
is thin and projects forward at an angle of 45°. The postero- 



medial wall extends backward to directly contact the 
pterosphenoid. 

The nasals (Fig. 4A) are large, almost entirely covering the 
lateral ethmoid and are narrowly separated from one another 
in the midline by the rostrodermosupraethmoid. Each bone 
has prominent anterior and posterolateral processes, two 
dorsal processes, lateral and medial, are folded inward to 
form curved flanges which support the skin roofing the 
sensory canal. In large specimens the nasals tend to become 
narrow with attrition of the anterior process (Fig. 6B). 

The frontals are nearly square except that the posterior half 
of the lateral border is deeply indented. Anteriorly, close to 
the midline is a high, arch-shaped crest (afc, Fig. 4A), a 
similar but longer arch is situated in the centre of the bone 
and is sometimes divided into two separate crests (cfc. 
Figs 4A, 5B), posteriorly is a low, diagonal crest (pfc, 
Fig. 4A). All these crests shelter a neuromast foramen and 
serve to support the skin covering the frontal canal system. 
Posteriorly, the frontal margin meets the pterotic, is over- 
lapped by the parietal and partially overlaps the anterior 
border of the supraoccipital. There are no ventral frontal 
laminae. The parietals (Figs 4A, 5B) are thin, near-diamond 
shaped bones each with a single neuromast foramen and 
posterolaterally covered by the median extrascapular. There 
is no parietal crest. 

The autosphenotic (Figs 4A,B, 5B) has a prominent, 
bluntly rounded lateral process and is overlapped by the 



ANATOMY OF THE MELANONIDAE 



17 



SO 



esc 




exo 



pro f/X bo v1 



Fig. 5. Neurocranium of M. zugmayeri in A, posterior and B, lateral views. In B, the intercalar is unshaded, the margins of the bones 
underlying it indicated by dashed lines. C, parasphenoid of 66mm SL specimen in dorsal view. 



frontal, parietal and pterotic. The underside of the bone 
bears a deep, almost transverse fossa into which articulates 
the hyomandibular. The pterotic (Figs 4A,B, 5A,B) accom- 
modates the posterior portion of the hyomandibular fossa 
along a third of its lateral border. The wall of the pterotic is 
somewhat bullate and its cranial surface forms a prominent 
lateral shelf. 

The pterosphenoid (Figs 4B, 5B) is long and deep forming 
most of the dorsomedial wall of the orbit, anteriorly it 
contacts the frontal and posteriorly the autosphenotic and 
prootic. The parasphenoid (Figs 4B, 5B,C) has a broad keel 
with, extending from its centre, a long, low ascending process 
which extends laterally at a low angle to the horizontal plane 
to meet the prootic; paired, parallel laminae rise from the 
central region of the keel to meet the bases of the lateral 
ethmoid wing (Fig. 6). 



The prootics (Figs 4B, 5B) are large with a deep trigeminal 
notch. The posterior border of the bone is rounded and 
partially overlapped by a relatively small, ovoid intercalar to 
which is attached the inferior limb of the 
posttemporal(Figs 4B, 5A,B, 6D). The small, pinnacle-like 
epioccipitals contact the posterolateral margins of the 
supraoccipital and posteriorly the dorsal borders of their 
respective exoccipitals; laterally each epioccipital is overlain 
by the second extrascapular (Fig. 4A). 

The exoccipitals are deeply depressed posteriorly and con- 
tain a large, backwardly facing vagus foramen (Figs 5A,B, 7). 
Medially, the bones meet across the midline by flange-like 
projections. Posteriorly there is an ovate, cartilage-filled 
process the base of which meets its antimere in the midline. 
Inside each exoccipital a long, ventrally directed process 
extends from the medial surface to contact a shallow dorsal 



18 



G.J. HOWES 




Fig. 6. Melanonus zugmayeri: A, vomer in ventral view; B, nasal 
of left side in dorsal view (broken outline indicates anterior nasal 
opening; C, extrascapulars of left side in lateral view; 
D, intercalar (left, lateral view). All from a specimen of 173mm 
SL. 

flange rising from the base of the basioccipital (Fig. 7B). The 
basioccipital is a trowel-shaped bone the blade of which forms 
the posterior basicranium and the handle, the occipital 
condyle (Figs 4,5,7). The supraoccipital (Figs 4A, 5A,B, 7) is 
well-ossified and lies flush with the frontals, its crest confined 
to its posterior margin; laterally, the bone is bevelled where it 
meets the parietal. Posteriorly its ventral margin is bordered 
by the exoccipital. 

The otoliths have been described and figured by Nolf & 
Steurbaut (1983; 1989). 

Comments on cranial features 

Melanonus has a plesiomorphic ethmo-vomerine region, 
namely a broadly rounded ethmoid lacking any dorsal eleva- 
tion as in macrouroids and with a single, narrow point of 
contact with the lateral ethmoid (Howes & Crimmen, 1990; a 
more extensive area of contact appears to be a feature of 
some supragadoids, Howes, 1990); a laterally expanded lat- 
eral ethmoid which contacts the ascending process of the first 
infraorbital ligamentously on its posterior face (Howes, 
1987); vomer with a relatively short shaft and well-formed 
teeth (absence of vomerine teeth in Macrouroidei and some 
gadoids is considered independently derived; see Okamura, 
1989; Inada, 1989; Howes, 1990). Ophidiiforms have as 
broad a variability of the ethmovomerine region as gadiforms 
but the lateral ethmoid is characterised by the presence of 
basal twin facets which firmly unite with the large palatine 
head. Furthermore, the lateral wing of the lateral ethmoid is 
usually reduced and feebly developed, but always has a 
lateral facet which articulates with the first infraorbital 
(Howes, 1992). 

The frontals of Melanonus have a plesiomorphic gadiform 
morphology; both gadoid and macrouroid taxa bear frontal 
crests of varying development as do ophidiiforms and this 



may be a 'paracanthopterygian' feature. Howes (1990:79) 
noted the lack of ventral frontal laminae in Melanonus and 
considered this a derived condition associated with the ante- 
rior displacement of the frontal area of the brain (see p. 27). 
Ventral frontal laminae are widely distributed amongst ophi- 
diiforms. There is no prominent V-shaped ridge pattern on 
the frontals in Melanonus and no 'mucosal' cavity, a feature 
of supragadoids. 

Nasal bones are plesiomorphically separated in the midline 
but in macrouroids are joined for most of their lengths, a 
feature regarded as synapomorphic for the group (Iwamoto, 
1989; Howes & Crimmen, 1990). Among gadiforms the size 
of the nasals is variable but they are nearly always large, 
trough-like bones containing two neuromasts. Among plesio- 
morphic gadoids (e.g. Bathygadidae) the size of the nasals 
approaches that of macrouroids but the bones remain sepa- 
rated along the midline. The melanonid condition is thus 
considered plesiomorphic although the nasal bones have a 
distinct apomorphic shape which more closely approaches 
that of some macrouroids than gadoids. 

The pterotic of Melanonus has a plesiomorphic gadiform 
morphology and resembles that of Bathygadidae in being 
broad with a rounded posterior margin and short hyomandib- 
ular fossa (Howes & Crimmen, 1990, fig. 6). 

The pterosphenoid is unusually large for a gadiform; the 
widespread condition (and among ophidiiforms) being small, 
occupying the dorsoposterior region of the orbit and widely 
separated from the lateral ethmoid. The enlarged anteriorly 
extended bone is therefore considered autapomorphic for 
Melanonus. The parasphenoid displays no particular derived 
feature and corresponds with the situation in the majority of 
gadiforms, namely a broad flat keel with parallel laminae 
(Howes, 1990:81). 

The deeply incised trigeminal notch of the prootic resem- 
bles most closely that of some Phycidae and the Muraenolepi- 
didae, but unlike those taxa the anterior wall of the prootic is 
directed medially as in most infragadoids and macrouroids 
(Howes, 1990:82). The intercalar is small in comparison with 
that in other gadiforms where in gadoids it is exceptionally 
large covering the entire posterolateral cranial wall. A large 
intercalar is one of the characters diagnostic of paracan- 
thopterygians but is secondarily absent in lophiiforms and 
batrachoidiiforms. Among ophidiiforms and percopsiforms 
the intercalar is generally not as large as that of gadiforms, 
and in those two former groups is confined to the upper half 
of the posterior cranial wall and does not extend ventral to 
the basioccipital anteriorly but is interrupted by the prootic. 

The relationship between the epioccipital and supraoccipi- 
tal is an unusual one amongst gadiforms in that the posterior 
walls of the exoccipitals meet across the midline and so 
exclude the supraoccipital from contributing to the upper 
margin of the foramen magnum. Elsewhere in paracan- 
thopterygians this condition occurs among ophidiiforms 
(Howes, 1992) where the exoccipital has enlarged backward 
and upward to cover the posterior margin of the supraoccipi- 
tal. Even in the largest cleared and stained specimen of M. 
zugmayeri examined for this feature the ventral tip of the 
supraoccipital does not reach the margin of the foramen 
magnum (Fig. 7B). The supraoccipital lacks a dorsal crest, 
there being a posterior lamina (Fig. 7). Howes (1990:82) 
discussed the variability of the supraoccipital crest amongst 
gadoids. An elevated cranial crest is possibly the plesiomor- 
phic condition for paracanthopterygians but a low, reduced 
crest is widely distributed amongst all groups and in lophii- 



ANATOMY OF THE MELANONIDAE 

so rial 



19 



exca 




boc 



Fig. 7. Melanonus zugmayeri posterior part of cranium in lateral oblique view of 130mm SL specimen showing in A, first neural arch and 
vertebra attached and in B, removed to expose the posterior features of the basi- and exoccipitals. Note the supraoccipital does not 
contribute to the border of the foramen magnum. 



forms and ophidiiforms appears to be the common condition. 
It is assumed that this feature has been repetitively evolved in 
these groups. 

Jaws (Fig. 8) 

The premaxilla (Fig. 8C) has tall, thin and widely separated 
ascending and articular processes, and a tall, spine-like 
postmaxillary process. The toothed surface is narrow, bearing 
for most of its length two rows of sharp pointed teeth. The 
outer row teeth are straight or extend slightly laterad, the 
inner row teeth which are about twice the length of the outer 
are inwardly curved; posteriorly there are three rows of teeth, 
the ones of the centre row being the same size as those of the 
inner (Figs 8D,E). In a 100mm SL specimen of M. zugmayeri 
the posterior teeth are so arranged as to form distinct 
transverse rows (Fig. 8F) but this is not evident in the 66mm 
or 130mm SL specimens. The maxilla has a tall articular head 
and a short medial articular process forming a rather acute 
angle with the head (Figs 8A,B). The shaft of the bone is 
slender and posteriorly bears shallow dorsal and ventral 
processes. 

The dentary (Fig. 8G) is short and deep with a correspond- 
ing shallow mentomeckelian cavity; it has a high steep 
coronoid process. The sharp pointed teeth are set in an 



irregular single row, numbering 22 in 88mm and 100mm SL 
specimens of M. zugmayeri, 2% in a 130mm and 34 in a 175mm 
SL specimen. The anterior teeth are small followed by four or 
five successively larger ones, then four or five relatively large 
teeth separated by three or four smaller ones. Posteriorly the 
teeth diminish in size. The anguloarticular (Fig. 8G) is tall 
with a steep posterior slope and short, vertical anterior 
margin; the articular condyle is long and narrow. The coro- 
nomeckelian bone (Fig. 8G) is a well-developed, cylindrical 
element with slight dorsal and ventral posterior flanges. The 
retroarticular (Fig. 8G) is boot-shaped, the leg being curved 
forward and the foot long and shallow. A strong labial 
ligament is anchored to the rim of the dentary (Howes, 1988, 
fig- 12). 

The overall jaw morphology of Melanonus is plesiomorphic 
for gadiforms, the upper jaw bones, apart from having a 
smaller postmaxillary process of the premaxilla, are little 
different from those in bathygadids (Howes & Crimmen, 
1990). Macrouroids are characterised by having a large 
postmaxillary process of the premaxilla situated posteriorly 
(Okamura, 1970; Howes & Crimmen, 1990). There is no 
'gadoid notch' at the base of the postmaxillary process. The 
lower jaw more closely resembles that of gadoids or ophidii- 
forms than macrouroids in having a relatively shallow angu- 
loarticular and boot-shaped or J-shaped retroarticular. 



20 



G.J. HOWES 











i^riWM^ , ^iHim'twr'iTr T « T * 



cm 




mc 



Fig. 8. Melanonus zugmayeri, jaw bones. A and B right maxilla from a 130mm SL specimen in: (A) dorsal and (B) medial and slightly 
ventral views; C, premaxilla in lateral view; D-F premaxilla, anterior (D) and posterior (E) regions from 130mm SL specimen and (F) 
100mm SL specimen, ventral views; G, lower jaw of 130mm SL specimen in medial view. 



Macrouroids tend toward a deeper anguloarticular and 
greater variability in the shape of the retroarticular (Oka- 
mura, 1970; Howes & Crimmen, 1990). A boot-shaped 
retroarticular is lacking in both percopsiforms and lophii- 
forms. 

Palatopterygoquadrate (Fig. 9) 

The palatine (Fig. 9A,B) is long, its posterior tip extending to 
nearly halfway along the ectopterygoid, its rostral process is 
long and slender and overlies the maxilla, its base bears a 
broad facet which articulates with the ethmoid cavity and the 
body of the bone rises to a high posterior crest. There are two 
rows of sharply pointed teeth. 

The anterior part of the ectopterygoid (Fig. 9 A) lies along 
the medial face of the palatine and its ventral stem reaches 
the quadrate joint; laterally it is slightly overlapped by the 
entopterygoid (Fig. 9A). The latter is a relatively large bone 
with a rounded dorsal profile and is sloped mesad, its 
posterior border is well separated from the hyomandibular by 
the metapterygoid. The metapterygoid (Fig. 9 A) is axe- 
shaped its posterior margin rising high up the leading edge of 
the hyomandibular shaft. 



The melanonid palatine is unique amongst gadiforms, in its 
length, nature of contact with the pterygoids, and in bearing 
teeth. The common condition, and one which is considered 
synapomorphic for gadiforms (p. 29) is for the palatine to be 
reduced in length with a vertical or slightly angled posterior 
border meeting a similar blunt margin of the ectopterygoid 
and forming a hinge-type joint (see figures in Okamura, 1970, 
1989; Howes, 1990, 1991b; Howes & Crimmen, 1990). This 
union differs from that commonly encountered in other 
paracanthopterygians where the posterior limb of the pala- 
tine is attenuated and articulates firmly with the leading edge 
of the entopterygoid and lateral face of the ectopterygoid. 
Percopsids resemble gadiforms in having a near vertical 
abutment of the palatine with the ento- and ectopterygoids 
(Fig. 9D). However, there is a posterior stem which overlaps 
the upper lateral margin of the ectopterygoid. Macrouroids 
are characterised by the lack of direct contact between the 
palatine and ethmovomerine bloc (Howes & Crimmen, 
1990). 

Norman (1930) noted there were 'teeth on the pterygoid 1 , 
an error perpetuated by Howes (1991b, caption to fig. 35). 
The pterygoid bones of Melanonus display plesiomorphic 
morphologies; the large entopterygoid and high posterior 



ANATOMY OF THE MELANONIDAE 



21 



— sop 




Fig. 9. A-C Melanonus zugmayeri: A, palatoquadrate, hyosymplectic and opercular bones of 130mm specimen in medial view, light hatched 
area represents ligamentous system connecting opercular bones to hyomandibular; B, palatine of 100mm specimen, right side, lateral view; 
C, hyomandibular, left side, of 100mm SL specimen in anterior view showing foramen for hyoid branch of facial nerve (arrowed) and 
lateral flange; D, Percopsis omiscomayus , palatine and pterygoids in lateral view (heavy dotting indicates cartilage). 



metapterygoid process are present in macrouroids, bathyga- 
dids and macruronids (Howes & Crimmen, 1990; Howes, 
1991b). Reduction of the entopterygoid and metapterygoid 
appears to be characteristic of supragadoids (Howes, 1990). 
Amongst ophidiiforms the metapterygoid abuts against the 
lower limb of the extended anterior portion of the hyoman- 
dibular. 

The quadrate (Fig. 9A) of Melanonus has a wide angle 
between its posterior border and the posteroventral spine. 
The size of this angle is variable among gadiforms and 
appears correlated with the orientation of the suspensorium. 
An 'interosseuos space' between the symplectic and preoper- 
culum (Okamura, 1970; 1989) is also a condition of the 
angular separation of the two parts of the quadrate, being 
absent where the angle is small (Howes, 1990). 



Hyoid arch (Figs. 9-11) 

The hyomandibular (Fig. 9A) has, as in all gadiforms, a single 
articulatory condyle. The bone is narrow-waisted with the 
relatively long shaft oval in section, a foramen for the hyoid 
branch of the facial nerve pierces its posterior margin (Fig. 
9C). Posteriorly is a long, horizontal process which articulates 
with the opercle. The lateroposterior face contacts the border 
of the preopercle. Medially a band of ligamentous connective 
tissue joins the shaft with the opercular process and a wider 
band runs at right angles to it to attach to the subopercle and 
interopercular-subopercular ligament (Fig. 9A). 

The course of the hyoid branch of the facial nerve is 
partially exposed laterally, due to attrition of the outer part of 
the hyomandibular, part of which remains as a lateral flange 
which is a common feature (synapomorph) for gadiforms, 



22 



G.J. HOWES 




Fig. 10. Melanonus zugmayeri hyoid bar of 100mm SL specimen: 
A, medial view; B and C, urohyal in lateral and dorsal views. 

one not shared by ophidiiforms or lophiiforms (Howes, 
1992). 

Other hyoid arch bones are much like those of the majority 
of gadiforms; the posterior half of the anterohyal =ceratohyal 
auct. (Figs 10A, 11 A) is deep and in this respect resembles 
that bone in some macrouroids (eg. Nezumia, Abyssicola, 
Coelorhynchus, Coryphaenoides; Okamura, 1970), more 
closely than gadoids. However, this feature is variable and a 
similar range of morphotypes can be found among ophidii- 
forms (Markle & Olney, 1990, fig. 13). As in most gadiforms 
and ophidiiforms there are 7 branchiostegal rays which 
appears to be the plesiomorphic paracanthopterygian num- 
ber, (six occur frequently in lophiiforms). The urohyal 
(Figs 10B,C) bears a closer resemblance to that of gadoids 
rather than macrouroids in having a shallow dorsal keel and a 
long, prominent anterodorsal (basibranchial) process 
(Howes, 1990, fig. 16B). 

The basihyal (Fig. 11 A) is a dumbbell-shaped bone lying 
between the dorsohyals and crossed by a ligament which 
connects them; anteriorly a thick cartilaginous 'tongue' pro- 
trudes forward, posteriorly, the basihyal is slightly over- 
lapped by the first basibranchial(see below). The interhyal 
(Fig. 9A) is typically gadiform, being long and slender, 
contacting the symplectic cartilage dorsally and the posterior 
socket of the posterohyal ventrally. Markle (1989, fig. 6A) 
shows a common ligamentous connection between the 
interhyal-posterohyal and interopercle. I find this to be one 
involving thick connective tissue although a discrete ligament 
runs from the medial side of the interhyal to the medial 
posterior tip of the posterohyal. 

Opercular bones (Fig. 9A) 

The opercular bones are relatively generalised except that the 
suboperculum has a straight to concave leading edge rather 
than the common gadiform condition of a rounded to pointed 
margin. The interopercle is shallow and nearly oblong with 
rounded dorsoposterior and anteroventral borders; it is 
widely separated from, and ligamentously connected to the 
subopercle. In general, macrouroids have the interopercle 
orientated horizontally (e.g. Okamura, 1970, figs 26; 27) 
whereas in gadoids the bone is angled, sometimes steeply as 
in Melanonus. Melanonus lacks the interopercular fossa 
present in a subgroup of 'supragadoids' (Howes, 1990). The 
opercle is relatively large for a gadiform and overlaps most of 
the subopercle. The preopercle has a short lower, anteriorly 
directed limb and a narrow laminate (symplectic) process 



which, plesiomorphically, contacts the symplectic cartilage. 
In its derived form the symplectic process of the preopercle 
contacts the lateral face of the hyomandibular (Howes, 1990). 

Branchial arches (Fig. 11) 

There are three basibranchials (Fig. 11 A), the first and 
second ossified, the third cartilaginous. The posterior margin 
of the first overlies the the anterior border of the diamond- 
shaped second which is separated from the small diamond- 
shaped third. The first and second hypobranchials (Fig. 11 A) 
are long with marked posterior curvature, both contacting the 
first basibranchial and bear gill-rakers on their outer and 
inner margins; the third is short and lacks gill-rakers. The first 
and second ceratobranchials (Fig. 11 A) bear five or six 
clustered-spinous rakers on their outer and the same number 
of slender, triple-spine rakers on their inner margins; the 
third has seven outer and inner shorter rakers and the fourth 
has four short rakers on its outer margin only. The anterior 
tips of the fifth ceratobranchials are apposed but not firmly 
united in the midline and are ligamentously connected to the 
third basibranchial; a narrow tooth patch bears ca 25 slender 
pointed teeth. 

The epibranchials (Fig. 11B) are 30% the length of the 
ceratobranchials. A strong uncinate process on the first 
epibranchial is connected by a chondrified ligament to a large 
interarcual cartilage; the third epibranchial bears a long tooth 
plate bearing ca 20 sharp pointed teeth. There are four 
pharyngobranchials (Fig. 11B), the first being an ossified 
element; the second-third pharyngobranchial tooth plates 
bear strong, pointed teeth. 

Markle (1989) has described and commented on the upper 




bM-3 



pbb1-3 



Fig. 11. Melanonus zugmayeri branchial arches of 130mm SL 
specimen: A, dorsal view of lower arch elements; B, ventral view 
of upper arch elements. In A, basihyal is also shown in lateral 
view. 



ANATOMY OF THE MELANONIDAE 



23 



branchial arch of Melanonus which he considers, due to the 
presence of a large and chondrified interarcual ligament, to 
be plesiomorphic for gadiforms. In his cladogram, however, 
he mistakenly ascribes to Melanonus the loss of the second 
pharyngobranchial. The lower gill-arch is also plesiomorphic 
in that the basibranchials are unexpanded and there is no 
forward ventral elongation of the third hypobranchial as in 
macrouroids and ophidiiforms; the first hypobranchial is 
typically long as in gadiforms but lacks an expansion where 
the ligament running to the dorsohyal attaches. Spinous 
gill-rakers of both the clustered- and triple-spine type are 
widespread amongst 'infragadoids' and macrouroids. 

Pectoral girdle (Fig. 12) 

The vertical and horizontal limbs of the cleithrum (Fig. 12A) 
are nearly equal in length; the medial cleithral lamina is thin 
and only prominent near the cleithral tip. Markle (1989) 
noted that the foramen which notches medial borders of both 
the scapula and coracoid is present only in the former 
(Fig. 12A). Markle (1989) and Howes & Crimmen (1990) 
commented on the variability of this feature; plesiomorphi- 
cally the foramen lies entirely within the scapula, a condition 



almost entirely confined to 'infragadoids' although it is also 
recorded in the 'supragadoid' Lota. The supracleithrum 
(Fig. 12B) is a lanceolate bone with a slightly expanded 
dorsal articulatory surface which contacts the posttemporal. 
There are four actinosts and 12 or 13 pectoral rays in the M. 
zugmayeri specimens examined (Norman, 1930, gives 13 for 
M.zugmayeri and 12-14 for M. gracilis; Fahay & Markle, 
1984, give a range for the genus of 10-16). The single 
postcleithrum (Fig. 12A) has a broad head and slender, 
slightly upwardly curved stem. It articulates in a cleft oppo- 
site or slightly above the coracoid-scapula junction (see also 
Markle, 1989, fig. 10). 

The posttemporal (Fig. 12B) is V-shaped, its upper limb 
broad proximally and tapering distally; its lower limb, which 
is firmly united with the intercalar is thin, rod-like and 
completely ossified. 

The extrascapulars (Figs 4A, 5A,B, 6C) number four, each 
having upturned borders and containing a neuromast. The 
lateral extrascapular covers the posterior corner of the 
pterotic, two lie in contact with one another along the medial 
part of that bone and the innermost lies along the lateral part 
of the parietal. In large specimens of M. zugmayeri the 




ac 





wt 




Fig. 12. Melanonus zugmayeri: A, pectoral girdle in medial view; B, posttemporal and supracleithrum in lateral views; C, pelvic girdle in 
dorsal view. 



24 



G.J. HOWES 



medial extrascapular is more closey aligned with the supraoc- 
cipital, resting along a lateral ridge of the bone and in a 
specimen of 173mm SL (Fig. 6C), it appears that the lateral 
and a medial extrascapular have become fused, judging by 
the presence of two neuromast foramina in the single large 
bone. 



Pelvic girdle (Fig. 12C) 

The pelvic bone is narrow and tubular, broadening proxi- 
mally where its cartilaginous tip contacts its antimere sym- 
physially. Distally the pelvic process is narrow and straight 
and connected with its antimere by ligamentous tissue. There 
is no lateral pelvic process or spine (cf. Bathygadidae, Howes 
& Crimmen, 1990). There are usually 7 fin rays; Fahay & 
Markle (1984) give a range for the genus of 5-7. 
The pelvic girdle lies well forward with the anterior tips of 



the pelvic bones lying between the cleithra so that the origin 
of the pelvic fin lies beneath or just anterior to that of the 
pectoral (Fig. 1). The position of the pelvic girdle in relation 
to the pectoral girdle is variable amongst gadiforms. In the 
majority of gadoids the pelvic girdle is situated well forward, 
particularly so in the more derived 'supragadoid' taxa such as 
gadids, gaidropsarids and muraenolepidids, so that the origin 
of the pelvic fin lies in advance of that of the pectoral fin. 
'Infragadoids' tend to have the pelvic girdle situated beneath 
or behind the pectoral (e.g. bathygadids, steindachneriids). 
In morids, however, the pelvic girdle lies well forward. There 
is some variability in position among macrouroids but gener- 
ally, the pelvic girdle lies posterior to the pectoral so that the 
origin of the pelvic fin is situated directly beneath that of the 
pectoral fin. With respect to the position of the pelvic fin 
relative to that of the pectoral, the Melanonidae appear to 
represent an intermediate condition between the derived 
forward and plesiomorphic posterior positions. 




Fig. 13. Melanonus zugmayeri vertebral column. A, first vertebra and neural arch of 130mm SL specimen in lateral and anterior views; B, 
anterior part of vertebral column of 100mm SL specimen showing retractor dorsalis muscle of one side, in ventral view; C and D vertebral 
column and fin supports of 130mm SL specimen: C, anterior vertebrae, D, 14th-17th vertebrae showing anterior anal fin supports (lateral 
views; ribs shown in black for clarity). 



ANATOMY OF THE MELANONIDAE 



25 



Vertebral column and median fins (Figs 13-15) 

There are 12-14 abdominal and 45 or 46 caudal (those with 
closed haemal spines) vertebrae in Melanonus (Fahay & 
Markle, 1984 give total counts of 58-62 for the genus). The 
first neural arch and spine are well-developed and form an 
ankylosed unit with the centrum. The prezygapophyses of the 
first vertebra (Fig. 13A) are oval in section, hollow and 
cartilage-filled and firmly in contact with the similarly shaped 
paired condyles of the exoccipital. The wall of the neural arch 
covers the upper posterior wall of the exoccipital leaving 
exposed a notch through which pass the occipital and lateral 
line nerves (Fig. 7B). The laminae of the neural arch extend 
forward to embrace the posterior extension of the supraoccip- 
ital crest (Fig. 7A). The second vertebra is anteroposteriorly 
compressed and lacks processes or ribs; the third-fifth verte- 
brae support successively shorter chopstick- shaped ribs 
which extend almost horizontally, at their tips lie epipleural 
(epineural) ribs the heads of which are ligamentously 
attached to their respective myosepta (Fig. 13B,C). The 
sixth-twelfth centra bear triangular parapophyses to each of 
which is attached a posteriorly curved epipleural rib. Accord- 
ing to Okamura (1989) there is a total of eleven epipleural 
ribs in Melanonus; ten are counted here in M. zugmayeri. 

There is a single dorsal fin comprising 72-78 rays. The first 
dorsal ray is often minute, the second and successive rays are 
long and flexible, supported by distally tapered rod-like 
radials which tend to occur in pairs within each interneural 
space, their proximal tips converging (Fig. 13C,D). The 
origin of the dorsal fin occurs between the third and fourth 
neural spines. There are no supraneurals (predorsals). The 
anal fin has 50-58 rays and lacks a stout anterior spine; the 
shape of the radials is similar to those which support the 
dorsal fin (Fig. 13D). 

Caudal fin skeleton (Fig. 15A,B). The caudal fin skeleton 
of Melanonus resembles that of the Moridae in that the first 
and second hypurals are incompletely fused; each support a 
single fin ray. In morids all the hypurals are fused only 
proximally whereas in Melanonus fusion of hypurals 1 and 2 is 
both proximal and distal leaving a central opening 
(Fig. 15A). Hypurals 3-5 although fused in specimens of M. 
zugmayeri of 130mm SL are only partially fused in 66mm and 
100mm SL specimens (Fig. 15B). Paulin (1983, fig. 5A) 
figures a caudal skeleton of M. gracilis in which hypurals 1 
and 2 are entirely fused and the fifth is reduced. In a 45mm 
SL specimen of M. gracilis, all the hypurals are separated for 
their entire lengths whereas in a 49mm SL specimen they are 
fused distally but not proximally. There are two elongate 
epurals each supporting a fin ray; in a 100mm SL specimen of 
M. zugmayeri they are joined proximally (Fig. 15B). Unlike 
morids, Melanonus lacks X and Y bones a feature shared with 
Macruronidae, Gadidae and Lotidae. A long parhypural 
articulates basally with the fused hypurals 1 and 2 and 
supports a single fin ray. 

Comments on features of the vertebral column and median 
fins. Chopstick-shaped ribs, similar to those of Melanonus, 
have been reported for Macruronus, Lyconus, Steindachneria 
and Merluccius by Okamura (1989) and Inada (1989) who 
arrive at opposite conclusions with regard to their character 
polarity. According to Okamura this rib-type suggests a close 
relationship between the taxa in which they occur. Inada, on 
the other hand, regards them as a plesiomorphic gadoid 
feature. Although Inada's (1989) reasoning appear to be 



based on an a priori assumption of merlucciid plesiomorphy I 
would agree with his assumption. In fact, this type of rib is 
more widely distributed amongst gadoids than has been 
reported and also occurs amongst 'supragadoids' other than 
Merlucciidae (Fig. 14A). 

In Melanonus the ribs occur on vertebrae 3-5 as in Lyco- 
nus, but they are on vertebrae 3 and 4 in Macruronus (both 
Macruronidae), 3-6 in Merlucciidae, and 3-4 in Gaidrop- 
saridae. In Steindachneriidae the ribs are on vertebrae 3 and 
4 but the rib on the fourth has less than half the thickness of 
that on the third whereas in the above cited taxa the ribs are 
of equal thickness. Furthermore, the epipleurals attach 
directly to the distal tips of the chopstick ribs in Steindachne- 
ria whereas in the other taxa they are indirectly attached by 
ligamentous strands running to the myosepta (as in Mel- 
anonus). In morids and 'supragadoids' epipleurals are 
attached directly to the vertebral ribs. Patterson & Rosen 
(1989) interpreted the vertebral ribs Steindachneria and 
Gadus as a parapophysis with attached epipleural. The ribs in 
Steindachneriidae, however, are like those of Melanonidae, 
Macruronidae and Merlucciidae in articulating with the ven- 
tral cavity of the centrum. In the Bregmacerotidae the third 
and subsequent vertebrae bear parapophyses to which are 
attached cartilage-formed ribs (Fig. 14B). The loss of epi- 
pleurals from the first and second centra is a gadiform 
synapomorphy (Markle, 1989). 

Howes (1991b) regarded the first neural arch of Macruro- 




epr 




Fig. 14. Anterior region of vertebral column in: A, Gaidropsarus 
mediterraneus; B, Bregmaceros sp. In A, black shading in the ribs 
(ar) indicates zones of cartilage. 



26 



G.J. HOWES 



P u2 puUul 




Fig. 15. Melanonus zugmayeri Caudal fin skeletons of A, 130mm SL and B, 100mm SL specimens. C, caudal fin musculature (although a 
superficial layer of connective tissue and some muscle has been removed the vertebrae are exposed in situ as shown). 



nus (Macruronidae) as a composite unit incorporating an 
accessory neural arch suggesting that the first centrum had 
been incorporated in the 'basioccipital'. Since, however, 1) 
ribs are always lacking from the first two vertebrae in 
gadiforms, 2) Baudelot's ligament always occurs on the first 
centrum and 3) an accessory neural arch does not occur above 
aulopiforms, it seems untenable that incorporation has 
occurred in macruronids. 

The caudal fin skeleton is lacking in the majority of 
gadiform taxa but where it does occur its most significant 
features are fusion of the upper hypurals and the presence of 
X and Y bones (lost in some taxa, see above), both of which 
contribute to the symmetry characteristic of 'supragadoids'. 
The morid caudal fin skeleton is regarded as the plesiomor- 
phic gadoid type since it approaches that of most other 
teleosts in its asymmetry and in having distally separated 
hypurals. In this latter respect, Melanonus demonstrates a 
further derived condition in having the hypurals distally fused 
(see further discussion on p. 30). 

Of particular note is the condition of the caudal fin 
musculature (Fig. 15C) which differs from that described in 
gadoids (Howes, 1991) where hypochordal longidorsales, 
flexores dorsales and inferiores are absent, the interradiales 
have a characteristic linkage pattern between the caudal fin 
rays and are continuous with the dorsal and anal fin rays. 
Howes (1991: 104) pointed out the absence of the latter in 
Melanonidae, but overlooked the fact that the caudal fin 
musculature more closely resembles that of other paracan- 
thopterygians and acanthopterygians in having discrete dorsal 



and ventral flexores and an amalgamated segment of interra- 
dialis musculature corresponding to the superficial interradia- 
lis. 

Melanonus, Lyconus and Brosme are the only gadiform 
taxa to possess a single dorsal fin, most have two and the 
more derived Gadidae have three. Dorsal fin origin is usually 
above the second and third neural spines, as in Melanonus, 
but the origin of the second dorsal is variable, the radial 
supporting the first ray of that fin being between the eight and 
ninth, ninth and tenth or tenth and eleventh neural spines. 
According to Inada (1989) the single dorsal fin of Lyconus 
evolved from amalgamation of two separate fins. Inada's 
evidence relies on a notch being present in the fin at a point 
above the proximally curved thirteenth radial which lies 
between the eighth and ninth neural spines which as just 
noted is the region commonly associated with the origin of 
the second dorsal fin. No similar 'evidence' occurs in Mel- 
anonus. 

Among gadoids the first radial of the first dorsal fin usually 
lies between the second and third neural spines, but this is 
variable being between the first and second in gaidropsarids, 
and in Bregmacerotidae the first radial has become directed 
forward so that the first dorsal fin ray lies above the supraoc- 
cipital. In macrouroids, the first supporting radial is also 
usually between the second and third neural spines but 
sometimes between the third and fourth. Percopsids, like 
melanonids have the first radial between the third and fourth 
neural spines. In ophidiiforms the position of the first radial is 
variable and can lie between any of the neural spines from the 



ANATOMY OF THE MELANONIDAE 



27 



first to the tenth. In batrachoidiiforms it is usually between 
the third and fourth neural spines and in lophiiforms the 
eighth and ninth or more posterior neural spines. 

Supraneurals, preceding the first dorsal fin are rarely 
present in gadiforms (Patterson & Rosen, 1989). 

Baudelot's ligament (Figs 13A,B) stems from the lateral 
cavity of the first vertebra to connect with the supracleithrum. 
The retractor dorsalis muscle originates from the fourth 
through sixth vertebrae; on the sixth it is attached to the 
leading edge of the parapophysis (Fig. 13B). 

Brain (Fig. 16). 

The brain of Melanonus is situated well forward, the telen- 
cephalon and anterior part of the mesencephalon being 
anteriorly displaced beyond the cranial cavity so as to lie in 
the orbital cavity formed by the enlarged pterosphenoids. 
The olfactory and optic lobes are large. The olfactory tracts 
are well separated and each tract is short and thick compris- 
ing at least twelve separate nerves each of which branches to 



feed the individual laminae of the nasal rosette. The olfactory 
bulb is large and lies against the lobe which is narrowly 
separated by a fissure from the laterally situated optic lobe. 
Upon leaving their respective lobes ventrally, the optic tracts 
cross and travel directly laterad a short distance to the eyeball 
which is only narrowly separated from the telencephalon. 
The cerebellar corpus is flat and lies pointing anteriorly 
between the optic lobes. This is a unique condition among 
gadiforms (noted by Marshall & Cohen, 1973 as diagnostic of 
the Melanonidae), normally the corpus is bulbous and ele- 
vated (Okamura, 1970) or lies posteriorly along the cerebel- 
lar crest. The cerebellar crest is flat and elongate flanked 
ventrolaterally by extensive trigeminal lobes. The cerebellar 
body extends posteriorly to entirely overlap the vagal lobes 
along the basal part of the medulla oblongata, also a unique 
gadiform condition. The granular eminence is large but not 
laterally extended. Ventrally, the inferior lobes, pineal body, 
hypophysis and vascular sac are all well-developed. 

The brains of some gadoids and macrouroids have been 
described by Svetovidov (1953), Okamura (1970) and Howes 



nr 




mo 



nVJI 



ge nV nau n ^ n % 



Fig. 16. Melanonus zugmayeri brain in A, dorsal and B, lateral views. In A, the pathways of the optic tracts beneath the lobes are indicated 
by dashed lines and the margin of the prootic is indicated by dashed lines lateral to the trigeminal-facial nerve complex. 



28 

& Crimmen (1990) and of those published descriptions Breg- 
maceros has the most similar overall morphology. Like that 
of Melanonus the brain is elongate with extensive trigeminal- 
facial lobes, a long cerebellar crest and closely connected 
olfactory bulb and lobe. However, there are major differ- 
ences in the relatively small size of the olfactory and inferior 
lobes and in the cerebellar corpus being orientated posteri- 
orly along the crest, having a posterolateral fissure and 
leaving the midline of the optic lobes exposed. 

Anterior placement of the forebrain was considered a 
gadiform character by Svetovidov (1948) and among gadoids 
there is a tendency for the brain to be shifted forward. In 
those few morids investigated and in macrouroids the fore- 
brain is generally confined to the cranial cavity. In some other 
paracanthopterygians (ophidiiforms, Howes, 1992 and per- 
copsids pers. obs.)the telencephalon lies in the orbital cavity 
as in Melanonus. 

It is problematic as to which features of gross brain 
morphology can be used as phylogenetic markers. The degree 
of separation of the olfactory bulb from the lobe is variable in 
gadiforms (discussed by Howes & Crimmen, 1990) but the 
plesiomorph condition, possessed by Melanonus, is seemingly 
for them to be closely associated. The shape of the olfactory 
lobe is also a highly variable feature and one that might, at 
least, be generically characteristic. 

Summarising data from gadoid brain descriptions given by 
Svetovidov (1953) it appears that elongate and short cerebral 
crests are equally distributed amongst the taxa he studied. A 
short, tall cerebral crest, common to gadoid brains, is also the 
common condition among paracanthopterygians. However, 
the granular eminence, although often large is laterally 
extended only in the Gadidae (sensu Dunn, 1989 and Howes, 
1991b). 



Swimbladder, viscera and body musculature 

(Fig. 17). 

The swimbladder is an elongate ellipsoidal, thin-walled sac 
adhering tightly to the vertebral column apart from where the 
long bilobed kidney runs on either side of the midline. 
According to Marshall & Cohen (1973) the melanonid swim- 
bladder is reduced and there are two retia. In the specimens 
of M. zugmayeri examined for this feature, the gas-gland 
covers nearly two-thirds of the anterior floor of the sac and 
there are four retia supplying separate lobes. Posteriorly the 
gas-gland tapers and is deeply pocketed. The oval appears to 
be beneath the retial area. 

The stomach is siphon-shaped, exceptionally thick-walled 
with a deeply and much convoluted mucosal membrane; 
there are six or seven caeca lying ventrally; the intestine is 
long and double-bended. The bilobed kidney is extensive, 
almost enveloping the stomach. The gonads lie posteriorly on 
either side of the swimbladder to which they are attached by 
thin strands. 

The anterior body musculature is similar to that described 
for Bathygadidae (Howes & Crimmen, 1990) except that 
melanonids lack the same degree of differentiation between 
dorsal and ventral sections of the epaxialis musculature (Fig. 
17), the dorsal section being apparent only anteriorly (the 
general condition) and not extended as far posteriorly as in 
bathygadids. 




Fig. 17. Melanonus zugmayeri anterior body musculature and 
visceral cavity dissected on the right side; the anterior part of the 
liver has been cut away to expose the pyloric caeca and the 
swimbladder has been dissected. 

DISCUSSION 



Melanonus has undoubtedly derived sensory features; the 
brain has a unique morphology amongst paracanthoptery- 
gians, extending well forward into the orbital cavity, the head 
is covered with a unique type and pattern of open-ended 
neuromasts innervated by the ramus canalis lateralis of the 
trigeminal nerve, the RLA nerve being absent. In its cranial 
osteological characters three can be considered derived: the 
shape of the fifth infraorbital, and exclusion of the supraoc- 
cipital from contributing to the foramen magnum. The first 
two of these osteological characters are autapomorphic; the 
enlarged pterosphenoid is undoubtedly correlated with the 
anterior position of the telencephalon. The third is a feature 
shared with ophidiiforms, in that group, however, the exoc- 
cipital is expanded dorsoposteriorly so as to exclude most of 
or the entire supraoccipital from the rear of the cranium and 
from contact with the first neural spine. In melanonids the 
supraoccipital is excluded from the border of the foramen 
magnum by its failure to extend ventrad during development, 
but nevertheless it still forms the upper posterior border of 
the cranium and contacts the first neural spine. 

Aside from these autapomorphies there are no other 
apomorphies which are shared with other gadoid taxa. The 
diagnosis of 'Gadoidei' has proved difficult since most syn- 
apomorphies so far proposed are either not exclusive to, or 
their distribution has not been completely documented, in the 
taxa currently embraced under this category (see below). 

Howes (1990; 1991a; 1991b) proposed a series of gadoid 
clades of which 'supragadoids' were recognised on the basis 
of a fused upper hypural plate of the caudal skeleton. A 
sequence of other synapomorphies, including an interopercu- 
lar fossa, contact of the posterior face of the lateral ethmoid 
wing by the first infraorbital and reduction of pterygoid bones 
(Howes, 1990) excluded Melanonidae from this group. Other 



ANATOMY OF THE MELANONIDAE 



29 



MELANONOIDEI MACROUROIDEI 



GADOIDEI 



^Bathygadidae Steindachneriidae Moridoidea Gadoidea] 




12. 13 



10:11 



Fig. 18. Proposed relationships of Melanonoidei with other gadiforms. Synapomorphies: 1, absence of pars jugularis, i.e. common aperture 
for principal cranial nerves (also occurs in some ophidiiforms); 2, loss of intermusculars from vertebrae 1 and 2; 3, scapular-coracoid 
foramen; 4, attrition of lateral face of hyomandibular; 5, levator arcus palatini covers lateral face of jaw musculature; 10, palatine forming a 
hinge or butt-joint with pterygoids; 11, enlarged intercalar contributing to posterior wall of cranium; 12, pharyngohyoideus muscle mediated 
by sternohyoideus; 13, interradiales muscle connected to dorsal and anal fin rays, loss of various caudal fin muscles and entire caudal skeleton 
in some taxa; 14, palatine contacts mesethmoid; 15, X and Y bones in caudal skeleton (lost in some laxa); 16, complete fusion of upper 
hypurals and symmetry of hypural plates. Autapomorphies for Melanonoidei; 6, supraoccipital excluded from margin of foramen magnum; 7, 
cranial neuromas! pattern and innervation; 8, brain position and morphology; 9, enlarged pterosphenoids contacting lateral ethmoids. 
Synapomorphies 1-5 and 10-11 from Gosline (1968; 1971); Howes (1988; 1989; 1990; 1991b); Markle (1989); Patterson & Rosen (1989). 
Synapomorphies for macrouroids summarized by Iwamoto (1989) and Howes & Crimmen (1990) and for moridoids by Paulin (1983). 



taxa so excluded are Moridae, Euclichthyidae, Steindachneri- 
idae and Bathygadidae. The two latter lack a caudal fin 
skeleton, thus the incomplete fusion patterns of hypural bones 
possessed by morids, euclichthyids and melanonids cannot be 
extended to these taxa. The cranial and vertebral osteology of 
Bathygadidae is plesiomorphic in comparison to other gadoid 
taxa whereas that of Steindachneriidae is relatively derived 
(pers. obs. see also Fahay's, 1989, notes on pelvic girdle 
morphology). 

The Melanonidae lacks a feature common to other 
gadiforms (macrouroids + gadoids), namely, a short palatine 
forming a butt or hinge joint with the ento- and ectopterygo- 
ids. In almost all gadiforms the palatine has a truncated near 
vertical margin which forms a mobile (laterally expanding) 
joint with the anterior margins of the pterygoid bones (p. 20). 
Melanonus has a plesiomorphic palatine where the stem 
firmly contacts the margin of the ectopterygoid. Moreover, 
the palatine extends some distance along the ectopterygoid 
and is toothed. Since no other gadiform has palatine teeth it 
might be assumed that the melanonid palatine is the primi- 
tively composite dermo- and autopalatine whereas other 
gadiforms have lost the dermal component. In other paracan- 
thopterygians, ophidiiforms and lophiiforms possess the ple- 
siomorphic, long posteriorly extended and toothed palatine; 



percopsids resemble gadiforms more closely in having an 
edentulous bone which abuts the straight anterior margins of 
the ecto- and entopterygoids but which still retains a posteri- 
orly directed stem (p. 20). 

The Melanonidae possesses three of those characters iden- 
tified by Patterson & Rosen (1989) and Markle (1989) as 
gadiform synapomorphies or potential synapomorphies, 
namely, absence of epipleural ribs from the first and second 
vertebrae; a scapular-coracoid foramen and absence of a 
lateral commissure, cranial nerves I— VII exiting through a 
common aperture. Two other potential synapomorphies 
listed by Patterson & Rosen (1989) are presence of X and Y 
bones and liver LDH pattern. X and Y bones are absent in 
melanonids and can only be judged as a plesiomorphic state 
or, against the congruence of other synapomorphies, as 
secondary loss. In the latter case the feature then appears as 
synapomorphic for a subgroup of gadoids (Fig. 18). LDH 
liver pattern has not been tested for in this taxon. 

Two other synapomorphies appear to be: 1) the form of the 
hyomandibular, which in the majority of gadoids and mac- 
rouroids has attrition of the anterior border and lateral face, 
fully or partly exposing the pathway of the hyoid branch of 
the facial nerve (Howes, 1989; 1991b; 1992); 2) the levator 
arcus palatini covering the adductor mandibulae musculature 



30 



G.J. HOWES 



laterally (Howes, 1988; 1991b). 

Melanonids have a small intercalar, a bone which in other 
gadiforms contributes to a substantial part of the lateroposte- 
rior cranial wall. In size the melanonid intercalar approaches 
that of Percopsis. Whether in Melanonus the bone is plesio- 
morphically small or whether there has been reduction sec- 
ondarily can only be assessed against the distribution of 
other, known derived features (Fig. 18). An intercalar is 
absent in lophiiforms and batrachoidiforms, an assumed 
secondary loss (Patterson & Rosen, 1989). 

The single dorsal fin is probably a plesiomorphic feature 
(p. 26). Among paracanthopterygians, an elongate second 
dorsal fin is assumed to be synapomorphic for anacanthines 
(sensu Patterson & Rosen, 1989). Melanonids share with 
ophidiiforms (including carapids and bythitoids), two gadoid 
genera (Lyconus and Brosme) and Macrouroididae a single 
dorsal fin which must be seen as resulting from either the 
'loss' of the first dorsal with anterior encroachment of the 
second, or the amalgamation of the two fins. It is impossible 
to distinguish between such phylogenetic events although 
either way the condition is seen as derived. Iwamoto (1989) 
considered the single dorsal fin of macrouroidids to be 
derived but that of the gadoid Brosme as plesiomorphic 
retention. Judging by the incongruent distribution of the 
character it is almost certainly homoplastic. The further 
partitioning into three fins in Gadidae represents a further 
derived state. 

In jaw musculature melanonids are little different from 
morids and bathygadids (Howes, 1988). Howes (1990; 1991b) 
noted a medial shift of adductor muscle Alb which would 
suggest a close phylogenetic relationship with supragadoids. 
This shift, however, is apparently induced by the presence of 
a unique transverse ligament which runs from the palatine to 
the inner face of the second infraorbital and which constricts 
and turns Alb inwards. This is not the same condition as the 
entire medial shift of an unconstricted Alb in 'supragadoids'. 

Melanonids have an unusual condition of the hyoid muscu- 
lature whereby the pharyngohyoideus (= rectus communis) 
attaches to the third hypobranchial as well as the urohyal 
(Howes, 1988). Urohyal attachment of the pharyngohyoideus 
is shared with macrouroids, two gadoid families and all other 
ctenosquamates (Lauder, 1983; Howes, 1988); in remaining 
gadoids the pharyngohyoideus is mediated by the sternohyoi- 
deus. It is assumed that the two exceptional gadoid families 
(Muraenolepididae and Ranicipitidae) have lost the sternohy- 
oideus attachment, the pharyngohyoideus being attached to 
the tip rather than the lateral face of the urohyal keel as it is 
plesiomorphically in melanonids. 

The melanonid caudal fin musculature (p. 26) lacks those 
features regarded as synapomorphic for gadoids (since mac- 
rouroids lack caudal fin skeletons and associated musculature 
it cannot be known whether this derived form of muscle 
arrangement was a gadiform feature subsequently lost in 
macrouroids). Melanonids have a caudal fin muscle arrange- 
ment only slightly modified from that present in other para- 
canthopterygians and in acanthopterygians. 

Although it cannot be doubted that the Melanonidae 
belongs among Gadiformes there is no evidence to suggest 
that it be regarded as a member of the Gadoidei. To be 
included within the Gadoidei, the elongate toothed palatine, 
lack of X and Y bones, reduced intercalar and single dorsal 
fin must be regarded as reversal and loss characters. The 
caudal fin skeleton demonstrates an advanced condition to 
that of the Moridae (the plesiomorphic gadoid taxon) in 



having, in adults, almost complete fusion of the upper 
hypurals which alone, would signify inclusion within the 
'supragadoids'. Indeed, I have argued elsewhere (Howes, 
1991b: caption to fig. 35) that the reported separation of 
hypurals in young ranicipitids (which I place amongst the 
'supragadoids') is a character reversal; a conclusion drawn on 
what appears to substantial support from other synapomor- 
phies. In the case of melanonids the principal evidence 
against the caudal fin skeleton being a character reversal is 
that the associated musculature has a plesiomorphic arrange- 
ment, lacking those derived elements found in the muscula- 
ture of morids and other gadoids, including ranicipitids (as an 
adult, Raniceps has the typically symmetrical gadoid caudal 
fin skeleton, lacking in Melanonus). Thus the fusion or partial 
fusion of the upper hypurals in melanonids is considered to 
have occurred independently to that in gadoids above the 
morid level. 

Taking into account these arguments and the anatomical 
evidence presented herein, the Melanonidae is regarded a 
basal gadiform taxon, representing, as Markle (1989) had 
previously hypothesised, the sister-group to both gadoids and 
macrouroids (Fig. 18). Such a phylogenetic arrangement 
leads to a higher level re-classification of the Melanonidae. 
Following Markle (1989) and recognising the family as being 
phylogenetically coordinate with the Macrouroidei and 
Gadoidei, it is placed in the suborder Melanonoidei. Those 
taxa which I have previously recognised as a monophyletic 
group termed 'supragadoids' are equivalent to Markle's 
(1989) Superfamily Gadoidea. The Moridae and Euclichthy- 
idae are regarded by Markle (1989) as sister taxa on the basis 
of asymmetry of procurrent caudal fin rays; I know of no 
supporting osteological synapomorphies for this relationship 
but provisionally accept it. Together these taxa form the 
sister-group to the Gadoidea and as such must be regarded as 
the Superfamily Moridoidea (= Moriformes, 

part,Schwarzhans, 1984). The 'infragadoids', Steindachneri- 
idae and Bathygadidae have no such status since they form an 
unresolved polychotomy with the Gadoidei + Moroidei and 
Macrouroidei. 



Acknowledgements. My thanks are due to Douglas Markle, Nigel 
Merrett and Colin Patterson for their helpful and critical comments 
on the manuscript of this paper. Since this is the final research project 
carried out while employed at The Natural History Museum I take 
this opportunity to thank all members of staff, associates and 
students past and present of the Fish Section for their advice, 
assistance, patience, generosity and friendship over the past twenty- 
five years. I am particularly indebted to my mentors, P. Humphry 
Greenwood and Ethelwynn Trewavas. 



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Bull. not. Hist. Mas. (Zool.) 59(1): 33-44 



Issued 24 June 1993 



A review of the serranochromine cichlid fish 
genera Pharyngochromis, Sargochromis y 
Serranochromis and Chetia (Teleostei: Labroidei) 

PETER HUMPHRY GREENWOOD 

Visiting Research Fellow, Zoology Department, The Natural History Museum, Cromwell Road, 
London SW7 5BD; Honorary Associate, J.L.B. Smith Institute of Ichthyology, Private Bag 1015, 
Grahamstown 6140, South Africa. 

CONTENTS 



Introduction 33 

Methods and Material 34 

Serranoch romine taxonomy 36 

Introduction 36 

The generic or subgeneric status of Serranochromis Regan, 1920, and Sargochromis Regan, 

1920, reconsidered 36 

The genus Chetia Trewavas, 1961 38 

The genus Pharyngochromis Greenwood , 1 979 39 

Conclusion 40 

The phyletic relationships of the serranochromines 40 

Generic key and diagnoses 41 

Acknowledgements 43 

References 43 



Synopsis. Recent taxonomic changes, newly described taxa and groups of taxa, and the introduction of taxonomic 
characters not previously employed require a revision of the informally recognised, essentially fluviatile group of 
southern African cichlids, the so-called serranochromines. The genera included in this assemblage are Pharyn- 
gochromis, Greenwood, Sargochromis Regan, Serranochromis Regan, and Chetia Trewavas. 

Previously, Sargochromis was considered to be a subgenus of Serranochromis, but new evidence indicates that it 
should be reinstated as a distinct lineage (i.e. genus). The species originally described as Serranochromis 
(Sargochomis) gracilis should now be transferred to the genus Chetia. Formerly the latter taxon was thought to be 
monotypic, but it is now expanded to include five species. One of these, Chetia brevis Jubb, had been included 
tentatively in the genus Astatotilapia, but is now returned to the genus in which it was described originally. 

The monophyletic origin of the serranochromines has still to be established. For that, and other reasons, doubt is 
cast on the phylogenetic reality of the 'Pharyngochromis - Chetia - Serranochromis' group of endemic species in 
Lake Malawi. The suggested interrelationships of the serranochromine genera presented below, and based on 
shared derived characters, cannot, for the same reasons, be considered a truly phylogenetic one. 



INTRODUCTION 



In a paper (Greenwood, 1979) reviewing and reconsidering 
the generic classification of several cichlid taxa then referred 
to the genus Haplochromis, an informal group of three 
genera was recognised on the basis of its constituent species 
having particular types of squamation and anal fin markings 
(op. cit: 229-316). The group was, and still is considered one 
of convenience because no cladistically based hypothesis 
could be erected to establish the monophyly of its contained 
genera, viz. Serranochromis Regan, 1920 (with which was 
incorporated, as a subgenus, Sargochromis Regan, 1920), 
Chetia Trewavas, 1961, and Pharyngochromis Greenwood, 
1979. A scheme of possible interrelationships of these taxa, 



suggested earlier by Trewavas (1964), was also discussed in 
my 1979 paper. 

Recent studies of the genera call for a revision of the 
group's taxonomy at the generic level, and a reconsideration 
of their possible interrelationship. For example, the genus 
Chetia, treated as monotypic by Greenwood, (1979) is now 
thought to contain five species (Balon & Stewart, 1983; 
Greenwood, 1984, 1992, and below, p. 38), certain problems 
regarding the generic classification of several Angolan hap- 
lochromine species have been clarified (Greenwood, 1979, 
1984 & 1992), new ideas on the supposed relationship of 
Serranochromis and Sargochromis have been put forward by 
Lippitsch (1991: 99-100), and Eccles & Trewavas (1989: 21) 
have formally recognised, amongst the endemic genera of 
Lake Malawi, a large assemblage of species which they 



34 



PH. GREENWOOD 



designate 'The Pharyngochromis - Chetia - Serranochromis 
group'. 

The term 'serranochromine' will be used in this paper as a 
group name for the genera Chetia, Serranochromis, Sar- 
gochromis and Pharyngochromis. Its use should not be 
construed as an indication or even a presumption of the 
group's ultimate and formal recognition as a Tribe. In the 
sense employed here it is comparable with my earlier use of 
the informal categories 'haplochromines' and 'pelmatochrom- 
ines' (Greenwood, 1979 & 1987). Such continued use, and 
introduction, of informal groupings clashes with the tribal 
status given by Poll (1986) to several cichlid assemblages in 
Lake Tanganyika, and with the geographically and taxonom- 
ically more extensive tribe Haplochromini defined by Eccles 
& Trewavas (1989), a tribe which also includes the serrano- 
chromines. In my view, these authors' actions are premature. 
Too few critical higher-level taxonomic studies have yet been 
made on the Cichlidae to support the phylogenetic relation- 
ships that are (or should be) implicit in the award of formal 
tribal status. For instance, Eccles & Trewavas (1989: 21) 
define the Haplochromini as: "Maternal mouth-brooding 
cichlid fishes of Africa and the Jordan Valley in which the 
basioccipital bone participates with the parasphenoid to form 
the apophysis for the upper pharyngeal bones". The value of 
the apophyseal character has been questioned by several 
authors (see review in Greenwood, 1978, also Greenwood 
1986) and it may have evolved more than once among 
African taxa (Greenwood, 1987); maternal mouthbrooding 
has apparently evolved independently in both the Tilapiini 
and Haplochromini (the tribes, respectively, sensu Trewavas, 
1983, and Eccles & Trewavas, 1989), and in one species of the 
genus Chromidotilapia of the pelmatochromines (sensu 
Greenwood, 1987: 169) in which paper it is also argued (op 
tit: 194-199) that this group should not be included, as it was 
by Trewavas (1983), in the tribe Tilapiini. 

Thus, the purpose of the present paper is simply to clarify 
the taxonomic status of the 'serranochromine' genera, and to 
establish a basis for further and phylogenetic studies of those 
taxa and those of Lake Malawi. 



METHODS AND MATERIALS 

Anal fin spots and true ocelli. One of the features used 
to define the serranochromines is the presence of maculae on 
the anal fin, usually in both sexes (Greenwood, 1979). A 
distinction was made there between true ocelli (such as occur, 
but almost exclusively in males, in a large number of hap- 
lochromine species, e.g. those in Lakes Victoria, Edward and 
Kivu), and the spots or maculae found in the serranochrom- 
ines and the haplochromines of Lake Malawi (see figure in 
Eccles & Trewavas, 1991). Judging by a recent description of 
a new Serranochromis species (Winemiller & Kelso- 
Winemiller, 1991) it is clear that some confusion still exists 
when discriminating between these two kinds of anal fin 
markings. Granted, it is often difficult to do so when only 
preserved material is available, but in life the difference is 
obvious, as colour photographs in aquarium books will show 
(e.g. Konings, 1991). 

The densely pigmented ovoid or near circular centre of the 
true ocellus, usually circumscribed by a narrow, darkly pig- 
mented ring, is surrounded, or almost surrounded, by a zone 
of virtually transparent, or at least freely translucent, and 



apparently unpigmented fin membrane. This clear zone is of 
variable width and outline, but is often concentric with that of 
the pigmented centre. In life, the clear zone seems to 
emphasise the coloured central area, thereby making it stand 
out from the rest of the fin membrane, be that membrane 
pigmented or hyaline. In colour photographs of live or freshly 
dead specimens, the clear zone often appears to be dark or 
even black, a result either of the dark background against 
which the fish was posed, or the shadow cast by the fish's 
body and the anal fin itself. To the best of my knowledge, 
true ocelli do not occur, at least in nature, on any of the other 
paired fins, although these fins are often maculate. Anal 
ocelli are also of rare occurrence in females, but large 
non-ocellate spots are sometimes present on that fin in the 
females of species whose males have true ocelli. The spots in 
such females occupy the same position as the ocelli in males, 
and are often of the same size. As compared with maculae 
(i.e. non-ocellate spots) on the anal fin, true ocelli are 
generally larger, and are always readily distinguishable from 
the maculae on the other unpaired fins. 

In contrast, non-ocellate anal spots, besides usually being 
smaller than ocelli, are, with few exceptions (see Greenwood, 
1992) more numerous and differ little in their overall appear- 
ance from those on the other fins, although the central 
pigmented portion may differ quite markedly in colour. The 
essential difference between maculae and ocelli, however, 
lies in the absence of a transparent or freely translucent area 
surrounding the pigmented centre of a macula, which, like 
that of an ocellus, is bounded by a very thin ring of dark 
pigment. Instead, the macula's pigmented centre is circum- 
scribed by a ring, usually narrow, of lighter pigment which 
separates it from whatever ground colour the fin membrane 
may have. 

This outer, lightly pigmented ring is not visible in some of 
the preserved serranochromine specimens I have examined, 
and the central spot is bounded only by the very narrow ring 
of dark pigment separating it from the chromatophores in the 
fin membrane. Whether or not this situation is a preservation 
artefact cannot be determined at present. 

In their account of the anal fin markings in the newly 
described species Serranochromis altus, Winemiller & Kelso- 
Winemiller (1991: 679) describe, unfortunately without an 
illustration, the fin in both sexes as having ". . .30-40 large 
pink or pink-orange ovate spots, each ringed with a transpar- 
ent, white ocellus. . .". I would suggest that the use of the 
words 'transparent and white' in apposition is somewhat 
contradictory, and that 'translucent white' would describe the 
condition more accurately, especially since it is one I have 
seen in fresh specimens of Serranochromis (and Sar- 
gochromis) species from the Okavango river and swamp 
system in Botswana. 

PRESHANK LENGTH OF THE MAXILLA; LENGTH OF THE PRE- 
MAXILLARY ASCENDING AND ALVEOLAR PROCESSES. The 

preshank length of the maxilla, relative to its shank length, 
and the length of the alveolar process of the premaxilla 
relative to the length of the entire ascending premaxillary 
process, are two morphometric characters not used in earlier 
papers (Greenwood, 1979, 1989, 1981). 

Preshank length of the maxilla is measured, on the bone's 
medial aspect, from the anterior tip of the medial arm of the 
maxilla's saddle process, to the mid-point of the anterior 
vertical projection from the upper margin of the bone's shank 
(see Fig. 1). Shank length is measured, also directly and on 



THE SERRANOCHROMINE GENERA REVIEWED 



35 



the bone's medial aspect, from the midpoint of the vertical 
projection to the posterior point on the maxilla's posterior 
margin (see Fig. 1). 




Fig. 1. Medial aspect of maxilla (from Serranochromis 
macrocephalus), viewed somewhat dorsolaterally, to show points 
of measurement for: A, preshank length, and B, shank length. 
Scale bar = 5 mm. 

The length of the premaxilla's ascending process is mea- 
sured, directly, from the dentigerous surface of the bone at its 
symphysis with the premaxilla of the opposite side, to the 
dorsal tip of the process. The length of the alveolar process 
uses the same ventral (i.e. dentigerous surface) reference 
point; its dorsal limit being the highest point on the dorsal 
margin of the process (see Fig. 2). 



STUDY MATERIAL 

See Greenwood (1979, 1984, 1992) for specimens used in 
previous studies of Serranochromis, Sargochromis, Chetia 
and Pharyngochromis . Additional material: 

Sargochromis coulteri RUSI 26627 '. Namibia (4 specimens) 
Sargochromis coulteri RUSI 36419. Namibia (1 specimen) 
Sargochromis coulteri RUSI 28106. Namibia (1 specimen) 
Sargochromis giardi RUSI 35782. Zambia (3 specimens) 
Sargochromis carlottae RUSI 31204 Namibia (3 specimens) 
Sargochromis carlottae RUSI 31214 Namibia (2 specimens) 
Sargochromis coulteri RUSI 31199. Namibia (1 specimen) 
Serranochromis robustus RUSI 31169. Botswana; Okavango 

river (1 specimen) 
Serranochromis thumbergi RUSI 22660. Dam, Empangeni 
area, Natal R.S.A. (1 specimen presumably an introduc- 
tion) 
Serranochromis longimanus RUSI 23877. Botswana; Oka- 
vango swamps (6 specimens) 
Serranochromis macrocephalus RUSI 24175. Botswana; Boro 

river (9 specimens) 
Serranochromis angusticeps RUSI 26854. Okavango swamps 

(4 specimens) 

Chetia brevis Holotype AMG. P. 951. Lomati river, R.S.A. 
Chetia brevis Paratypes AMG. P. 952. Lomati river, R.S.A. 

(5 specimens) 
Chetia brevis AMG. P. 1422. Lomati river, Barbeton district, 

R.S.A. (3 specimens, one partly skeletonized; see below) 
Chetia flaviventris AMG. P. 1298. Mogel river, Waterberg, 

R.S.A. (7 specimens, one partly skeletonized; see below) 
Chetia flaviventris AMG. P. 6871. Tweeport, Rustenburg 
district, Limpopo system, R.S.A. (10 specimens) 




Fig. 2. Frontal view of left and right premaxillae (from 
Serranochromis macrocephalus) to show points of measurement 
for: A, height of ascending process, B, height of alveolar process. 
Scale bar = 5 mm. 

Chetia flaviventris Palala river at Muisvogelkraal (24° 00'S, 

28° 24' 30"E) R.S.A. (5 specimens) 
Chetia mola Holotype. ROM 29825. Luonga river (Zaire 

drainage), Zambia 

Skeletal material 

Pharyngochromis acuticeps RUSI 36553. Okovango river 

(see also Greenwood, 1992) 
Sargochromis carlottae RUSI 31204. Namibia 
Sargochromis carlottae RUSI unregistered. 142 mm S.L. 
Sargochromis giardi RUSI unregistered 210 mm S.L. 
Sargochromis giardi RUSI unregistered 244 mm S.L. 
Sargochromis cfSargo. greenwoodi RUSI unregistered 
Serranochromis macrocephalus RUSI 24175. Botswana, Boro 

river 
Serranochromis macrocephalus RUSI unregistered 107 mm 

S.L. 
Serranochromis macrocephalus RUSI unregistered 220 mm 

S.L. 
Serranochromis macrocephalus RUSI unregistered ca 

220 mm S.L. 
Serranochromis angusticeps RUSI unregistered 220 mm S.L. 
Serranochromis angusticeps RUSI unregistered 230 mm S.L. 
Serranochromis angusticeps RUSI unregistered 410 mm S.L. 
Serranochromis longimanus RUSI unregistered 150 mm S.L. 
Chetia flaviventris AMG P. 1298. Mogel river, Waterberg, 

R.S.A. 
Chetia brevis AMG P. 1422. Lomati river, R.S.A. 

Institutional abbreviations: 



AMG 
ROM 
RUSI 



Albany Museum, Grahamstown 
Royal Ontario Museum, Toronto 
J.L.B. Smith Institute of Ichthyology, Graham- 
stown 



36 



P.H. GREENWOOD 



SERRANOCHROMINE TAXONOMY 



Introduction 

In their revision of certain haplochromine genera from Lake 
Malawi, Eccles & Trewavas (1991: 21) divide the non- 
tilapiine taxa of that lake into three groups, one of which they 
call the Pharyngochromis - Chetia - Serranochromis group. 
That action I consider to be premature, both because there is 
little concrete information available about the phyletic inter- 
and intrarelationships of the Malawi species, and because, as 
Eccles & Trewavas point out, there are differences between 
the squamation of those species and that of the serrano- 
chromines as construed in this paper (see p. 40). Granted, I 
have suggested (Greenwood, 1979: 314) that Serranochromis- 
and Chetia-like taxa could have been involved in the origin of 
the Malawian cichlid flocks, but that idea was not put forward 
on the basis of characters constituting a testable hypothesis. 
Rather, it was intended, because of the superficial resem- 
blance between the two groups, to promote an awareness that 
possible intergroup synapormophies should be looked for in 
future research. 

As recognised in this paper, the serranochromines are an 
assemblage of mainly fluviatile taxa whose geographical 
range encompasses the Zambezi, Save-Runde, Limpopo, 
Cunene, Quanza, Okavango and Zaire river systems, with 
one species (Serranochromis robustus robustus [Giinther]), 
occurring in Lake Malawi (see Balon & Stewart, 1983; 
Bell-Cross, 1975; Eccles & Trewavas, 1989; Greenwood, 
1984, 1992; Jubb, 1967, 1968; Ladiges, 1964; Poll, 1967; 
Skelton, 1993, Trewavas, 1961, 1964). 

Since the last published inventory of serranochromine 
species (Greenwood, 1979), revisional studies (Greenwood, 
1984, 1992) and the description of new species (Balon & 
Stewart, 1983, Winemiller & Kelso-Winemiller, 1991, and 
Greenwood, 1984 [see p. 38 below]) have both increased the 
number of included taxa and extended the geographical range 
of the group. 

Morphologically, the two distinguishing features of the 
serranochromines are the following, (i) The presence, often 
in both sexes, of non-ocellate maculae (see p. 34) on the anal 
fin. Generally these spots are very numerous with, in certain 
species, as many as 30-40 covering almost the entire fin. 
However, their number and size show considerable intraspe- 
cific (and intergeneric) variability, with as few as three or four 
large spots occurring in some individuals of a species where 
the maximum number is 18-20 (Greenwood, 1992). When 
many spots are present their arrangement may give the 
impression of an irregular distribution on the fin, but (as was 
noted by Oliver [1984], pace Greenwood, 1979: 315) there is 
a basically linear regularity in their arrangement, (ii) All 
scales above the lateral-line series are cycloid, as are the 
majority of scales below that level. Some weakly ctenoid 
scales may occur anteriorly on the flanks, especially in small 
specimens, the ctenii being confined to a narrow arc situated 
near the centre of the scale's free margin. 

A third but less trenchant feature of the serranochromines, 
as compared with other fluviatile non-tilapiine and non- 
pelmatochromine taxa (both sensu Greenwood, 1987: 
194-199) is a tendency for there to be a higher modal number 
of abdominal vertebrae (modes 16 or 17 in serranochromines, 
[but 14 in one taxon] cf 12 or 13 in the other taxa); however, 



the ranges for total vertebral counts in the two groups 
overlap. 

Where information is available on breeding habits, the 
serranochromine species are known to be female mouth- 
brooders, and in all taxa the neurocranial apophysis for the 
upper pharyngeal bones is formed from the parasphenoid and 
the basioccipital bones. In those respects, the group would 
conform with Eccles & Trewavas' (1989) definition of the 
tribe Haplochromini. 

Recently, Lippitsch (1991) drew attention to the possible 
value of scale morphology and patterns in resolving certain 
problems of intergeneric relationships within the serrano- 
chromines. To determine if the features described by Lip- 
pitsch could also provide another 'group' characteristic, I 
examined, at a magnification of 50x, the superfical morphol- 
ogy of flank and other scales in several species belonging to 
both subgenera of Serranochromis (sensu Greenwood 1979; 
but see p. 37), all species of Chetia, and in different popula- 
tions of the single Pharyngochromis species. In general I 
found a fairly high level of intrageneric variability, as well as 
some individual variability in the features noted by Lippitsch 
(1991: 99-100; figs D & E) i.e. ornamentation of the caudal 
field, and the presence of a soft caudal rim to the scale. 
Further studies will be necessary before the value of these 
features can be established, both at the group and lower 
levels of serranochromine taxonomy. However, another 
squamation feature noted and discussed by Lippitsch, namely 
the number of scale rows between the posterior orbital 
margin and the preoperculum, has proved of considerable 
value in reviewing generic level taxonomy within this group. 

The generic or subgeneric status of Serranochromis 
Regan, 1920, and Sargochromis Regan, 1920, 
reconsidered. 

In an attempt to revise the Haplochromis generic concept on 
phyletic lines, several so-called Haplochromis species were 
assigned, as a subgenus, to the genus Serranochromis (Green- 
wood, 1979). Since one of these species is the type of Regan's 
genus Sargochromis (S. codringtoni [Big, 1908]) that name 
was resurrected for the new subgenus. 

The principal argument for this taxonomic rearrangement 
was that the taxa included in the new concept of Serrano- 
chromis all share what appeared to be three derived features, 
viz: an increased number of abdominal vertebrae, higher 
gill-raker counts, and an increased number of branched rays 
in the dorsal fin (Greenwood, 1979: 299). 

In the light of new data, especially those stemming from an 
increased knowledge of the genus Chetia (see p. 38) and, 
particularly, Lippitsch's (1991) observations on the different 
type of postorbital squamation patterns present in the two 
presumed subgenera, I would now revise my earlier action 
and recognise both Serranochromis and Sargochromis as 
distinct lineages, and thus accord each generic status. The 
reasons for that action are as follows. 

Firstly, as Lippitsch (1991) noted, in Serranochromis but 
not in Sargochromis there are at least two, and often more, 
vertical rows of scales between the posterior orbital margin 
and the upper part of the preoperculum's ascending arm. In 
Sargochromis only a single row is present. A double row, 
however, also occurs in Chetia (personal observations, see 
below). Further investigation of both Serranchromis and 
Chetia reveals that the double and multiple scale row condi- 
tion is correlated with underlying osteological and myological 



THE SERRANOCHROMINE GENERA REVIEWED 



37 



characters that are not present in Sargochromis or Pharyn- 
gochromis, the two other serranochromines with only a single 
postorbital scale row. These correlated features are an 
increase in the relative size and bulk of the upper part of the 
levator arcus palatini muscle, and an anteriorly directed 
lengthening of the postorbital process from the sphenotic 
bone, particularly its ventral region associated with the origin 
of the muscle (see fig. 3 and fig. 8 in Greenwood, 1992). In 
Sargochromis and Pharyngochromis the muscle is relatively 
smaller, and the postorbital sphenotic process has the almost 
uniformly narrow form (Fig. 3) found in the generalised 
haplochromine skull (and, it may be added, the skulls of 
Tylochromis and Heterochromis , the genera thought to repre- 
sent the least derived lineages of African cichlids; see Oliver, 
1984, and Stiassny, 1989 & 1991). 

The other reason for reconsidering the subgeneric status of 
Sargochromis is the increased information now available, 
especially for Chetia (see p. 38), which clearly indicates an 
extensive overlap in the gill-raker and dorsal fin-ray charac- 
ters previously used (Greenwood, 1979) to define Serrano- 
chromis (then including Sargochromis). These characters, 
therefore, no longer can be considered synapomorphic for 
Serranochromis and Sargochromis alone. 

With the elimination of those two characters, the only 
derived feature shared uniquely by Serranochromis and Sar- 
gochromis is the increased number of abdominal vertebrae. 
Against that presumed synapomorphy must be set the 
derived postorbital scale-row character shared only by Serra- 
nochromis (sensu Regan, 1920) and Chetia, which genera also 
share two apomorphic features not previously recognised. 
These are: (i) an increase in the range and modal number of 
caudal vertebrae (range 15-18, modes 16 and 17 in Serrano- 
chromis, range 15-17 modes 16 and 17 in Chetia, compared 
with ranges of 12-16 in Sargochromis , and 14-16 in Pharyn- 
gochromis, with modes of 14 and 15, and 15 in the genera 
respectively); (ii) an increase in the range and modal number 
of circumpeduncular scale, viz. 18-20, rarely 16, in Serrano- 
chromis and Chetia, neither taxon with a clear-cut modal 
number, as compared with 16 or 18 (mode 16) in Sar- 
gochromis and 15 or 16 (mode 15) in Pharyngochromis . The 
recognition of this feature as an apomorphy is based on the 
circumpenduncular counts for Tylochromis and Hetero- 
chromis (see above), where the range, and modes, are 15-16. 

Thus, taking into account the presumed derived characters 
shared only by Serranochromis and Chetia, it would seem that 
the higher count of abdominal vertebrae in Serranochromis 
and Sargochromis should be treated as a homoplasy and not, 
as previously thought, an apomorphy indicating an immedi- 
ate common ancestor for the two taxa. For that reason I agree 
with, and now formally act upon Lippitsch's (1991) conclu- 
sion that "It seems advisable ... to recognise Sargochromis 
as a distinct genus. . .". 

The possible relationship of Sargochromis within the serra- 
nochromines is discussed on p. 41, and a revised generic 
diagnosis is given on p. 42. 

A few further comments need to be made about the genera 
Serranochromis and Sargochromis. The number of postor- 
bital scale rows in certain Serranochromis species can be as 
high as four or five, and, as far as I can determine, the 
number of rows is relatively constant intraspecifically. At 
present insufficient information is available on possible inter- 
or intraspecific differences in scale ornamentation, or on such 
variation in the presence or absence of a soft free margin to 
the scales (see Lippitsch, 1991). The small sample of species I 




Fig. 3. Posterior portion of the neurocranium, in left lateral view, 
to show differences in the size of the sphenotic postorbital process 
(Sp.) in Sargochromis, Chetia and Serranochromis. A, 
Sargochromis carlottae (RUSI: 31204, 89 mm S.L.; B, Chetia 
brevis (AMG: P1422, 90 mm S.L.) and C, Serranochromis 
macrocephalus (RUSI: 24175, 112 mm S.L.) Scale bar = 5 mm. 



38 



PH. GREENWOOD 



have examined from the Okavango swamp and river system 
in Botswana seems to indicate that such variation may occur. 
A soft margin, for example, was absent in specimens of 
Serranochromis thumbergi and S. longimanus, but is present 
in S. macrocephalus . From the same samples some individual 
variability was noted both in the extent of the granular area 
on the caudal field of flank scales below the upper lateral- 
line, and the presence or absence of a granular area in at least 
some scales above that line. Apparent interspecific variation 
is also seen in the extent and nature of cheek, opercular and 
interopercular squamation patterns. The value of these fea- 
tures as a basis for intrageneric classification has, however, 
yet to be tested on a wider geographical and taxonomic basis. 

A feature of Serranochromis, Sargochromis and Chetia 
noted in an earlier paper (Greenwood, 1979), where it was 
ranked as an apomorphy, is a reduction in the maximum 
number of inner tooth rows in both jaws to a single or 
irregularly double series. The new species, 5. altus, described 
by Winemiller & Kelso-Winemiller (1991), is exceptional in 
having as many as six rows anteriorly, reducing to a single or 
double row posteriorly (the figures adjusted from Winemiller 
& Kelso-Winemiller who include the outer tooth row in their 
counts). These authors also note that the number of inner 
rows increases with growth ". . .in all species of Serrano- 
chromis that we have examined. . .". Their list of study 
material indicates, however, that only one other species, S. 
angusticeps, was studied, and that the maximum number of 
inner or outer rows in that species is three. Since museum 
specimens are often not fully representative of a species' full 
size-range, the possibility of growth related changes in the 
number of tooth rows should perhaps be treated, for the 
moment, as an open question. In that connection, however, it 
should be noted that in Sargochromis, one species, Sargo. 
thysi, has four inner rows in both jaws at a size when only one 
or two rows are present in other congeneric species. 

Another dental feature in Serranochromis, one also shared 
with Sargochromis, (and probably Chetia), is for the two 
median teeth in the outer row of the inner tooth series to be 
enlarged and displaced anteriorly (Greenwood, 1984: 216; 
also see figs 3 & 8 in Trewavas, 1964). Amongst the sample of 
Serranochromis specimens examined, which included all spe- 
cies of the genus except S. altus. (see p. 35, and Greenwood, 
1979 & 1984 for additional material), such tooth displacement 
occurs in most species but not in all individuals of a species. 
Its frequency of occurence is lower in Sargochromis than in 
Serranochromis, and the condition is known from only one 
Chetia species (C. gracilis; see below and p. 39). Because of 
this inconstancy in its expression I would now consider the 
character more in the nature of a trend, albeit a derived one, 
rather than the trenchant synapomorphy recognised earlier 
(see Greenwood, 1984; 216). 

A revised generic diagnosis for Serranochromis is given on 
p. 42, and the possible relationships of the genus are dis- 
cussed on p. 40. To the list of included species published in 
Greenwood, 1979 (pp. 302-303) must be added 5. altus 
Winemiller & Kelso-Winemiller (1991) from the Zambezi 
system. 

Little need be added to previous accounts of the genus 
Sargochromis, type species Paratilapia codringtoni Blgr., 
1908 (see Greenwood, 1979; 1984, in both papers the taxon 
treated as a subgenus) except to note that the species 
described as Serranochromis (Sargochromis) gracilis Green- 
wood (1984) from the Cunene river, is now considered to be a 
member of the genus Chetia (see p. 39). The transfer of this 



species reduces to two the number of Sargochromis species 
having, relative to other congeneric taxa, slender lower 
pharyngeal bones with few and only partially molariform 
teeth, viz. Sargochromis greenwoodi (Bell-Cross) and Sarg. 
coulteri (Bell-Cross). The latter species, however, appears to 
show considerable variability in these features, with some 
individuals having noticeably coarser lower pharyngeal bones 
than do others, a feature invariably correlated with an 
increased number of molariform teeth (Greenwood, 1984: 
217-221). As was discussed in that paper, the species level 
taxonomy of Sargochromis is far from satisfactory, so this 
seemingly intraspecific variability should only be accepted 
with some reservation (see also Greenwood, 1965 and 
Hoogerhoud, 1986). 

In lacking a pronounced ventral expansion of its sphenotic 
postorbital process(see p. 37), the neurocranium in Sar- 
gochromis is immediately distinguishable from that of Serra- 
nochromis (see Fig 3; also comments in Greenwood, 1979: 
303, and figs 13 & 16). 

A list of Sargochromis species is given in Greenwood, 
(1979, pp. 304-305), and a revised generic diagnosis on p. 42 
below. 

The genus Chetia Trewavas, 1961 

Type species. Chetia fla viventris Trewavas, 1961. 

It has not proved possible to formulate a trenchant generic 
definition for Chetia since the taxon has yet to yield a single 
diagnostic autapomorphy. Its definition, therefore, is based 
on, as it were, negative features, namely those which exclude 
the five member species from inclusion in any of the three 
other serranochromine genera, especially Serranochromis 
with which Chetia has the greatest superficial and some 
detailed similarity. Thus, although Chetia shares with Serra- 
nochromis the apomorphy of two postorbital scale rows (see 
p. 36), a high number of caudal vertebrae (15-17, modes 16 
or 17) and an increased number of circumpeduncular scales 
(18 or 20, rarely 16), it does not share with Serranochromis 
apomorphies of an increased number of abdominal verte- 
brae, (and as a consequence, an increase in the total number 
of vertebrae), nor the increased lateral-line scale count of 
that genus (35-41, a character probably correlated with the 
increased number of vertebrae). Neither do the two genera 
share the autapomorphic dental character of Serranochromis , 
namely the development of a totally unicupsid jaw dentition 
at a very small size; i.e. , at some length, yet to be determined, 
less than 29 mm S.L.; see Greenwood, (1979: 300). 

On the basis of that analysis, the simplest diagnosis for 
Chetia is: a Serranochromis lacking the apomorphic features 
of that genus (see also Greenwood, 1992: 49-50 and p. 43 
below). 

At the time of my earlier generic review (Greenwood, 
1979), Chetia comprised two species, namely the type, Chetia 
flaviventris , and Chetia brevis, Jubb, 1968; the species coming 
from the Limpopo and Incomati river systems of South Africa 
respectively. 

Based on Jubb's original description together with an 
examination of the holotype and two other Chetia brevis 
specimens, I excluded, in my 1979 paper, the species from 
Chetia and placed it, as incertae sedis, in the genus Astatotila- 
pia (Greenwood, 1979: 284 & 307). That decision was based 
on three supposed features of C. brevis. (i) Jubb's (1968) 
description of the anal fin markings as ocelli, together with 



THE SERRANOCHROMINE GENERA REVIEWED 



39 



his and my personal observations of their large size and 
restricted number (3 or 4) as compared with the small and 
numerous anal spots in other serranochromines (see p. 34), 
especially Chetia flaviventris. (ii) Jubb's (op. cit) statement 
that the scales are ctenoid, and its implication that such scales 
are the predominant form, (iii) The presence of bicuspid 
teeth in the outer oral tooth rows of some specimens more 
than 100mm S.L., contrasting with the situation in Chetia 
flaviventris where, on the information then available, few 
bicuspids were thought to be present in fishes 71-85 mm 
S.L., and only unicuspids occurred in specimens above that 
length. 

Having now been able to examine colour-transparencies of 
live C. brevis, it is clear that the anal spots are not true ocelli 
(as defined on p. 34), but are large versions of the maculae 
found in other serranochromines, including Chetia flaviven- 
tris. The taxonomic significance to be attached to their large 
size, and small number, cannot yet be assessed. However, a 
wide size-range of anal maculae, their size negatively corre- 
lated with their number, occurs in the monotypic serrano- 
chromine genus Pharyngochromis , some individuals of which 
have spots as large as those in Chetia brevis. 

The larger collection of C. brevis specimens now available, 
together with a re-examination of the entire type series, 
shows that Jubb's blanket description of the scales as ctenoid 
is somewhat misleading. As in Chetia flaviventris , the scales 
of C. brevis are cycloid above the lateral-line, and mostly 
cycloid below that level. A variable number, usually small, of 
ctenoid scales is present anteriorly on the flanks, such scales 
being most numerous in fishes less than 80 mm S.L. In other 
words, the same pattern as occurs in Chetia flaviventris is also 
found in C. brevis. 

An examination of larger samples of both C. flaviventris 
and C. brevis also revealed that bicuspid teeth can be present 
in idividuals of C. flaviventris up to a standard length of 
90 mm, and that true unicuspid teeth as well as very weakly 
shouldered (almost unicuspid) teeth occur in C. brevis speci- 
mens between 80 and 90 mm S.L. This situation severely 
weakens my third reason (see above) for excluding C. brevis 
from the genus Chetia. 

Furthermore, the presence of two vertical rows of postor- 
bital scales in both Chetia brevis and C. flaviventris (see 
p. 36), a feature previously overlooked by all workers, pro- 
vides additional evidence for returning the species 'brevis' to 
the genus in which it was originally, and I now acknowledge 
correctly placed by Jubb (1968). 

Since 1979 another two species have been included in the 
genus, namely the taxon Chetia mola described by Balon & 
Stewart (1983) from the Zaire river system, and Hap- 
lochromis welwitschi (Blgr.) from Angola (see Greenwood, 
1979 & 1984). 

Chetia mola differs from all other congeneric species in 
having a greatly enlarged lower pharyngeal bone with a 
heavily molarized dentition (see fig. 12 in Balon & Stewart 
1983), a type of pharyngeal mill more usually associated with 
species of Sargochromis . However, unlike members of that 
genus, C. mola has a double or sometimes triple row of 
postorbital scales, and a lower number of abdominal verte- 
brae. 

To the existing four species of Chetia, I would now add a 
fifth, a species from the Cutato river, Angola, originally 
described (Greenwood, 1984) as Serranochromis (Sar- 
gochromis) gracilis. The reasons for this transfer are the 
double row of postorbital scales in 'gracilis', the persistence 



of compressed, bi- and weakly bicuspid teeth in the outer 
tooth row of both jaws in specimens over 100 mm S.L. and of 
tricuspid teeth in the inner tooth rows in such invididuals (see 
Greenwood, 1984: 227, and above), and the presence of only 
15 abdominal vertebrae, a combination of derived and plesio- 
morphic characters not occurring in any known Sargochromis 
or Serranochromis species. 

On the basis of comparisons with the few specimens 
available, Chetia gracilis (n=2) differs from the only other 
Chetia species recorded from southwestern Africa, C. wel- 
witschi (n=5), in having a narrower interorbital width 
(18.2-18.6 cf 21.0-23.3% of head length in C. welwitschi), a 
larger eye (25.0-25.6, cf 18.6-22.25% head; the effects of 
allometry are unlikely to account for this difference since the 
two samples overlap in the size range of individuals repre- 
sented), and a shallower cheek (22.7-23.3 cf 32.0-33.3% 
head). The lower pharyngeal bone in C. gracilis is somewhat 
stouter, and its median teeth coarser than in C. welwitschi (cf 
figs 19 & 20 in Greenwood, 1984). 

In its morphometric and meristic features, C. gracilis 
closely resembles C. flaviventris and C. brevis, species respec- 
tively from the Limpopo and Incomati river systems in South 
Africa. From Chetia brevis, C. gracilis is distinguished by its 
narrower interorbital width (18.2-18.6 cf 23.0-26.0% head 
length), and from C. flaviventris, especially when specimens 
of approximately the same size are compared, by its shal- 
lower cheek (22.7-23.3 cf 25.5-33.6% head). It also seems 
likely that, when the effects of allometric growth are taken 
into account, the eye diameter is greater in C. gracilis than in 
C. flaviventris. The teeth in the median row of the lower 
pharyngeal bone of C. gracilis are coarser and stouter than 
those of C. flaviventris, in that respect being more like the 
teeth in C. brevis (cf fig. 19 in Greenwood, 1984, fig. 19 in 
Greenwood, 1974, and fig. 4B in Jubb, 1968). 

Regrettably, apart from Chetia flaviventris (see du Plessis 
& Groenewald, 1953; Trewavas, 1961) and C. mola (see 
Balon & Stewart, 1983; fig. 10b) little or nothing is known 
about the live coloration of the other Chetia species. 

A revised generic diagnosis for Chetia is given on p. 43. 

Included species: 

Chetia flaviventris Trewavas, 1961. Limpopo river system. 

Chetia brevis Jubb, 1968. Incomati river system 

Chetia mola Balon & Stewart, 1983. Luongo river, Zaire 

system. 
Chetia welwitschi (Boulenger), 1898. Cunene and Zaire river 

drainage systems, Angola (see Greenwood, 1979). 
Chetia gracilis (Greenwood), 1979. Cutato river (Cubango 

drainage system), Angola. 

The genus Pharyngochromis Greenwood, 1979. 
Type SPECIES: Pelmatochromis darlingi Boulenger, 1911. 

A detailed revision of this monotypic genus, together with 
an annotated synonymy, has been published recently (Green- 
wood, 1992). Three nominal species all classified in the genus 
Haplochromis since Regan's revision of 1922 (or in one case, 
Pharyngochromis) , viz. Pelmatochromis darlingi Boulenger, 
1911; Chromis jallae Boulenger, 1896, and Pelmatochromis 
multiocellatus Boulenger, 1913, are now treated as junior 
synonyms of the single Pharyngochromis species, P. acuticeps 
(Steindachner) 1866. 

There are some indications that P. acuticeps could be 



40 



P.H. GREENWOOD 



considered either as a superspecies composed of several 
topospecies or as an assemblage of evolutionary species, but 
no clear-cut features allowing a formal taxonomic division of 
the taxon can be identified (see Discussion in Greenwood, 
1992). 

Anatomically and morphologically, Pharyngochromis is 
the least derived member of the serranochromines. Its sole 
autapomorphic feature is the higher position occupied by the 
posterior scales of the upper lateral-line series relative to the 
base of the dorsal fin (see Greenwood, 1992). The last five to 
seven (rarely four or eight) pored scales in that series are 
separated from the dorsal fin base by only one large and one 
much smaller scale. In the other serranochromine genera 
only the last one or two pored scales are separated from the 
fin base in this way, the other posterior scales having at least 
two large scales of equal size interposed between them and 
the fin base (see Greenwood, 1979 & 1992). 

In most P. acuticeps specimens (from all localities) the 
lower pharyngeal bone is slightly enlarged, and some of its 
median row teeth, which are always coarser than their lateral 
congeners, have molariform or submolariform crowns 
(Greenwood, 1992, fig. 7). There is, however, considerable 
individual variability in the degree to which this bone is 
enlarged and its teeth are molarized. In these respects, P. 
acuticeps resembles certain Sargochromis species, particularly 
S. coulteri (Bell-Cross) and S. greenwoodi (Bell-Cross). But, 
since the two genera differ in several other features, I would 
consider this resemblance to be homoplastic and not, as 
Trewavas (1961: 9) suggested, one of phylogenetic signifi- 
cance. 

If the serranochromines are a monophyletic lineage, (see 
p. 41) their recent common ancestor could well have resem- 
bled the extant Pharyngochromis acuticeps, except for the 
incipient hypertrophy of the pharyngeal jaws in the latter. 

Pharyngochromis acuticeps has a very wide distribution 
which includes the Zambezi and Save-Runde river systems, 
the Okavango river and its delta swamps, Lake Calundo, the 
Lucala river (Quanza drainage) and some unidentifiable 
localities in Angola. Records of the species (as Haplochromis 
darlingi) from the Limpopo river system (Jubb, 1967, 
repeated in Greenwood, 1979) are now known to be errone- 
ous and probably based on the misidentification of small 
Chetia flaviventris specimens. 

A revised generic diagnosis of Pharyngochromis is given in 
Greenwood (1992:48) and on p. 42 below. 



CONCLUSION 

The phyletic relationships of the 
serranochromines 

The difficulties encountered in determining both the inter- 
and intrarelationships of these fishes were discussed in two 
previous papers (Greenwood, 1979 & 1992), as was Trewa- 
vas' earlier (1964) tentative scheme of their relationships. 
Basically the problem lay, and still lies, in establishing 
whether or not the group is of monophyletic origin. 

One of the two features unifying the serranochromines, the 
non-ocellate anal fin markings, is probably a plesiomorphic 
character, perhaps representative of a stage in the evolution 
of true ocelli and one in which the markings, unlike true 
ocelli, are not confined to males (Greenwood, 1979; Oliver, 



1984). However, the possible function of anal maculae as 
egg-dummies (sensu Wickler, 1962; 1963), or even if they 
play any part in reproductive behaviour (Hert, 1989), have 
yet to be determined. There is also the possibility that 
non-ocellar anal markings have evolved more than once 
within the haplochromine cichlids (sensu law), as their pres- 
ence in species of Thoracochromis (Greenwood, 1979, 1984) 
could suggest. (Alternatively, Thoracochromis and the serra- 
nochromines may be more closely related than other morpho- 
logical and anatomical evidence would indicate.) In this 
context, Eccles & Trewavas', (1989: 27) obervations on three 
species of the endemic Malawi genus Aulonocara are perti- 
nent. One of these species has simple spots, a second has 
spots surrounded by a contrasting border, and the third no 
markings at all, although the fin has a pale border. Since 
Aulonocara has a number of distinctive anatomical autapo- 
morphies, this situation certainly suggests that not only have 
different kinds of anal fin markings evolved more than once 
but have done so within a single genus. 

Another aspect of the problem involving phylogeny and 
anal markings is Oliver's (1984: 108) suggestion that hap- 
lochromines with multiple non-ocellar spots, together with 
those having true ocelli, comprise a monophyletic assemblage 
whose members are more closely related to one another than 
to any species with what he considers to be the pleisiomorphic 
condition of anal maculae, namely spots indistinguishable 
from those on the other unpaired fins, especially the dorsal 
fin. That hypothesis has yet to be tested by the identification 
of suitable congruent and derived features characterizing the 
two supposed lineages, and raises questions about the signifi- 
cance of the seemingly unique anal marking of Pseudo- 
crenilabrus species (Greenwood, 1989). 

All in all, the current evidence for anal fin markings being 
of value in reconstructing phylogenies is not encouraging, 
particularly at the taxonomic levels under consideration here. 
For that reason I cannot agree with Eccles & Trewavas' 
(1989) use of the feature as grounds for suggesting a close 
relationship between the fluviatile serranochromines and 
most of the endemic haplochromines of Lake Malawi. While 
agreeing that non-ocellar anal spots are plesiomorphic fea- 
tures, these authors believe that the absence of true ocelli in 
the two groups ". . .combined with the geomorphological 
history of the region . . . may be accepted as evidence for the 
relationship of the Malawian group of species with the 
haplochromines of the Zambezi area" {i.e., with Serrano- 
chromis, Sargochromis and Pharyngochromis; Chetia has not 
been recorded from the Zambesi system). Certainly the 
similarity in anal fin markings would seem to support the 
intuitive feeling that the two groups could be related (see 
p. 34), and thus encourage a search for other characters to 
confirm or refute that impression, but in itself I would not 
rate it as 'evidence'. 

The second morphological character used to define the 
serranochromines, i.e. cycloid scales above the lateral line 
and a preponderance of such scales below that level, is, I 
would argue, a derived condition (Greenwood, 1979; see also 
Oliver, 1984; Lippitsch, 1991). Nevertheless, it is not clearly 
an autapomorphic feature of the serranochromines. For 
example, in the Lake Malawi haplochromines mentioned 
above, there is also a marked reduction in scale ctenoidy, but 
here, although scales above the lateral-line, like those in the 
serranochromines, are cycloid, the ctenoid scales occurring 
below that level are confined to the posterior part of the body 
and not the anterior part as in serranochromines. In a 



THE SERRANOCHROMINE GENERA REVIEWED 



41 



phylogenetic context how are these similarities and differ- 
ences, to be evaluated? 

Again, a pattern of reduced ctenoidy like that in the 
serranochromines occurs in some but not all species of 
Thoracochromis (Greenwood, 1979; 291; 1984:192 & 200), a 
genus in which, apparently, there are both true ocellar and 
non-ocellate types of anal fin markings (see above), but with 
both kinds occurring only in males. Possibly the 'genus' 
Thoracochromis is polyphyletic and that some of its species 
should be included in the serranochromine assemblage. 
Here, as is so often the case, one is hampered both by a 
paucity of detailed information on live coloration and the 
relatively few specimens available for anatomical and mor- 
phological studies. 

Another difficulty, linked with lack of information, lies in 
the possibility that further research could establish that there 
really is a close phylogenetic relationship between the serra- 
nochromines and certain haplochromines of Lake Malawi. In 
that eventuality, it is possible that the nearest relative of one 
or more of the fluviatile serranochromine taxa is to be found 
in the lake's fauna, thus rendering the serranochromines, as 
currently conceived, either a para- or a polyphyletic group. 

Although at present the monophyly of the serranochrom- 
ines cannot be established or refuted, it is possible to con- 
struct, on the basis of shared derived features, a tentative 
intragroup taxonomy. 

As compared with Pharyngochromis , the genera Chetia, 
Serranochromis and Sargochromis all share two derived fea- 
tures (see Liem, 1991) associated with the upper jaw skele- 
ton, viz an increase in the shank length of the maxilla relative 
to its pre-shank length, and an increase in the length of the 
alveolar process of the premaxilla relative to the length of the 
entire ascending process of that bone (see Methods). In 
Pharyngochromis the preshank portion of the maxilla is from 
1.2-1.3 times longer than its shank length, whereas in Chetia 
and Sargochromis the two parts are of equal length (with, in 
some Sargochromis species the preshank portion slightly 
shorter) and in Serranochromis the shank is noticeably longer 
(as much as 1.3 times so). In Pharyngochromis the length of 
the alveolar process of the premaxilla's ascending process is 
60-66% of the length of the entire ascending process; in 
Sargochromis it is from 69-76%, in Chetia 13-11% and in 
Serranochromis 73-83%. 

Neither of these ratios, either inter- or intragenerically, 
appears to be influenced by the size of the 17 specimens 
examined, all in the size range 79-410 mm S.L. and repre- 
senting ten species. 

On the basis of those two characters, a Pharyngochromis 
and a Chetia - Serranochromis - Sargochromis subgroup can 
be recognised within the serranochromines. Both these 
derived features are the only ones shared by the three latter 
genera (the increased number of abdominal vertebrae used 
previously to unite Sargochromis and Serranochromis in a 
single genus [Greenwood, 1979] is now thought to be a 
homoplasy; see p. 37). 

Chetia and Serranochromis both share three derived fea- 
tures not found in Sargochromis or in Pharyngochromis viz. 
(i) an increased modal number of caudal vertebrae (range 
15-17, modes 16-17 in Chetia, and 15-18, modes 16 and 17 in 
Serranochromis) compared with Sargochromis (range 12-16, 
modes 14 and 15) and Pharyngochromis (range 14-16, mode 
15). (ii) Two or more vertical rows of postorbital scales (c/a 
single row in Sargochromis and Pharyngochromis , see p. 36). 
(hi) An increased number of scale rows around the caudal 



peduncle, i.e. 18-20, rarely 16, compared with 16-18 rarely 
15, (mode 16), in Sargochromis, and 15 or 16 (mode 15) in 
Pharyngochromis . 

On the basis of those three derived characters, and using 
the term 'sister taxon' without any phylogenetic implications, 
then, within the serranochromines, Pharyngochromis is the 
sister taxon to the group Sargochromis, Chetia and Serrano- 
chromis, and within that latter group, Sargochromis is the 
sister taxon of Serranochromis and Chetia combined. 

That scheme bears, in broad outline, a close resemblance 
to Trewavas's (1964) diagram suggesting the interrelation- 
ships of Serranochromis, a scheme based essentially on 
lateral-line and dorsal fin ray counts (which may, of course, 
be correlated, in part, with the vertebral counts used here) 
and certain characteristics of the pharyngeal jaws. 

When comparing the two schemes, allowances must be 
made for the fact that two of Trewavas' Haplochromis species 
(lucullae and darlingi) are now treated as synonyms of 
Pharyngochromis acuticeps (Greenwood, 1992), that the sta- 
tus of H. angolensis, H. humilis and H. toddi is uncertain or 
unknown (Bell-Cross, 1975, Greenwood, 1979), and that H. 
welwitschi is now considered to be a species of Chetia (see 
p. 39). Also, the two Haplochromis species, mellandi and 
frederici, placed in limbo between the genera Haplochromis 
and Sargochromis in Trewavas' diagram, are now included in 
Sargochromis (Bell-Cross, 1975; Greenwood, 1979 and above 
p. 37), as is Haplochromis carlottae. 

Trewavas (op. cit. 9-10) superimposed on her diagram a tree 
indicating the possible phylogenetic relationships of the taxa, 
both inter- and intragenerically. It is here that our views would 
not coincide, mainly because I do not think there is the evidence, 
based on cladistic methodology, to justify the relationships 
proposed, even at an intergeneric level (see p. 40). Certainly 
there are no features justifying Trewavas' (op. cit. 10) suggestion 
that Serranochromis is a diphyletic and gradal taxon or that a 
cladal grouping would recognise Chetia, Serranochromis robus- 
tiis and S. thumbergi on the one hand, and Chetia welwitschi 
(Trewavas' Haplochromis welwitschi), and the remaining Serra- 
nochromis species on the other. Nor can I accept Trewavas' 
uncertainty about the separation of Sargochromis codringtoni, 
mellandi, carlottae and greenwoodi (as Haplochromis frederici in 
her scheme; see Bell-Cross, 1975) from the 'Haplochromis' (i.e. 
Pharyngochromis,) root of her tree (see Greenwood, 1979, 1992 
and above). A truly phylogenetic assessment of the groups' 
relationships must, as discussed on p. 36, await the results from 
further and preferably multidisciplinary research into the system- 
atics of all the serranochromine species, and those many Malawi 
species that Eccles & Trewavas (1989: 21) have placed in their 
Pharyngochromis - Chetia - Serranochromis group. 

Generic key and diagnoses 

A single row of scales between the posterior orbital margin 

and the vertical limb of the preoperculum A 

Two or more scale rows between the posterior orbital margin 
and the vertical limb of the preoperculum B 

A. Last 2 or 3 pored scales in the upper lateral-line series 
separated from the dorsal fin base by not less than two scales 

of approximately equal size A(i) Sargochromis 

Last 5 to 7 (rarely 4 or 8) pored scales in the upper lateral line 
separated from the dorsal fin base by one large and one small 
scale A(ii) Pharyngochromis 



42 



PH. GREENWOOD 



A(i): 

Abdominal vertebrae [15] 16-18 [19], modes 16 and 17; 
caudal vertebrae 12-16, modes 14 and 15; total number of 
vertebrae 28-32 (mode 31). 

Dorsal fin with 13-16, modes 15 and 16, spinous rays and 
11-16, modes 12 and 13, branched rays. Anal fin with 3 spines 
and 8-11, mode 9, branched rays. Caudal fin truncate, 
subtruncate or almost rounded. 

Scales in the lateral series 28-34, modes 30 and 31. Cheek 
with 3-6 horizontal rows. [15] 16-18, mode 16, scales around 
the caudal peduncle. 

Gill-rakers in the outer series on the first ceratobranchial 
9-15, modes 12 and 13. 

Outer series of teeth in both jaws composed mainly of 
unequally bicuspid teeth in fishes <150 mm S.L., but pre- 
dominantly of unicuspids in larger specimens. Inner series of 
jaw teeth, except in one species, arranged in a single or 
double series anteriorly and anterolaterally, reducing to a 
single row posterolaterally; in the exceptional species there 
are 4 rows anteriorly and anterolaterally, and a single row 
posterolaterally. 

Pre-shank length of the maxilla equal to, or slightly shorter 
than the shank-length (see p. 34). Height of the premaxillary 
alveolar process 69-76% of the height of the entire ascending 
process (see p. 35). For comments on neurocranial morphol- 
ogy and other osteological features (including the lower 
pharyngeal bone and its dentition, see text and figures in 
Bell-Cross (1975) and Greenwood (1979: 303-305, figs. 
16-18; and 1984: 216-225, figs. 12-17). 

Lower pharyngeal bone hypertrophied in the majority of 
species, greatly so in some, but only slightly enlarged in two 
species. The extent and degree to which the dentition of this 
bone is molarized is positively correlated with the degree of 
the bone's hypertrophy. In species with slightly enlarged 
bones only a few molar-like or submolariform teeth are 
present, and are confined to the median tooth rows. The 
ventral outline of the bone's anterior keel is almost straight 
and rarely extends below a horizontal drawn through the 
deepest point on the bone's ventral surface below the denti- 
gerous area. In specimens with a greatly enlarged lower 
pharyngeal bone, however, the ventral margin of the keel 
extends a little below that level (see figs 12-17 in Greenwood, 
1984). 

Anal fin spots small and numerous (as many as 40) 

Sargochromis 

A(ii): 

Abdominal vertebrae [13] 14 or 15 [16], mode 14; caudal 
vertebrae 14-16, mode 15; total number of vertebrae 28-31, 
mode 30. 

Dorsal fin with 14-16, mode 15, spinous rays and [10] 
11-13 [14] branched rays. Anal fin with 3 spines and 8 or 9 
branched rays (no distinct modal number). Caudal fin dis- 
tinctly truncate, subtruncate or almost rounded. 

Scales in the lateral series [30] 31-36, mode 33, modal 
range 32-34. Cheek with [3] 4-6 horizontal rows. 15, rarely 
16, scales around the caudal peduncle. 

Gill-rakers in the outer row on the first ceratobranchial 
7-12, modes 9 and 10. 

Outer series of jaw teeth composed of unequally bicuspid 
teeth in fishes <80 mm S.L., although some unicuspids can 
be found in larger fishes within that length range. Unicuspids 
become the predominant form in fishes >90 mm S.L. Inner 
series of teeth, in both jaws, arranged in 1-3 rows anteriorly 



and anterolaterally, reducing to a single row posterolaterally. 
The number of inner rows anteriorly appears to be positively 
correlated with an individual's size. 

Pre-shank length of the maxilla clearly greater than the 
shank length (see p. 34), i.e. about 1.2-1.3 times longer. 
Height of premaxillary alveolar process 60-66% of the height 
of the entire ascending process. For comments on neurocra- 
nial form and other osteological features, see Greenwood 
(1992). 

Lower pharyngeal bone in most individuals showing a 
slight degree of hypertrophy. In specimens over 50 mm S.L., 
the median rows of lower pharyngeal teeth are composed of 
noticeably coarser teeth than those situated laterally, and 
some can have submolariform crowns; the degree of molar- 
ization is most marked in fishes over 100 mm S.L. Irrespec- 
tive of the degree to which the lower pharyngeal bone is 
enlarged, its anterior keel is deep, with a curved ventral 
outline whose deepest point lies below a horizontal drawn 
though the deepest point of the ventral surface underlying the 
dentigerous part of the bone (cf Sargochromis above); see fig. 
7, Greenwood, 1992. 
Anal fin spots of variable size and number, from as few as 3 

or 4 large spots to as many as 19 small ones 

Pharyngochromis. 

B. 16-18 (rarely 15 or 19) abdominal vertebrae; inner and 
outer rows of jaw teeth composed entirely or mostly of 
unicuspids in fishes over 30mm S.L. (and posibly in smaller 

individuals as well) B(i) Serranochromis 

14 or 15 abdominal vertebrae; many bicuspid (or weakly 
bicuspid) teeth present in the outer tooth rows of both jaws in 
fishes as large as 80 mm S.L B(ii) Chetia 

B(i) 

Abdominal vertebrae [15] 16-18 [19], modes 16 and 17; 
caudal vertebrae [15] 16-18, modes 16 and 17; total number 
of vertebrae 31-36 (no distinct modes). 

Dorsal fin with 13-18, modes 15 and 16, spinous rays, and 
13-16, modes 14,15 and 16, branched rays. Anal fin with 3 
spines and 9-13, modes 10 and 11, branched rays. Caudal fin 
subtruncate or almost rounded. 

Scales in the lateral series [34] 35^41, no distinct modes. 
Cheek with 3 (rarely) to 11 horizontal rows (modally 5-9 
rows). 18-20 scales around the caudal peduncle (no distinct 
mode). 

Gill-rakers in the outer series on the first ceratobranchial 
[8] 9-13, modes 10,11 and 12. 

Outer and inner series of jaw teeth composed of unicuspids 
in specimens over 30 mm S.L. Inner series of both jaws, in all 
but one species, arranged in a single or double row (rarely 3 
rows) anteriorly and anterolaterally, and a single row poster- 
olaterally. In the exceptional species there are as many as six 
rows anteriorly and anterolaterally, reducing to a single or 
double row posterolaterally. 

Pre-shank length of the maxilla shorter than its shank- 
length (see p. 34), which is ca 1.2-1.3 times longer than the 
pre-shank portion. Height of the premaxillary alveolar pro- 
cess 73-82% of the height of the entire ascending process (see 
p. 34). For comments on the neurocranium and other osteo- 
logical features see Greenwood (1979: 299-302; figs. 13-15) 
and Trewavas (1964). 

Lower pharyngeal bone slender, its dentigerous surface 
elongate and narrow (see figures in Trewavas, 1964, and 
Greenwood, 1979). No molariform pharyngeal teeth; even 



THE SERRANOCHROMINE GENERA REVIEWED 



43 



those teeth in the median rows are only a little coarser than 
the other and fine teeth on the bone. 

Anal fin spots small and numerous (as many as 40) 

Serranochromis 

B(ii): 

Abdominal vertebrae 14 or 15 (no distinct mode); caudal 
vertebrae 15-17, modes 16 and 17; total number of vertebrae 
30-32, mode 31. 

Dorsal fin with 14 or 15, mode 15, spinous rays and 10-13, 
modal range 11 or 12, branched rays. Anal fin with 3 spines 
and 7-10 branched rays (no distinct mode). Caudal fin 
truncate to subtruncate. 

Scales in the lateral series 32-35, modal range 32-34. 
Cheek with 4-6, modes 5 and 6, horizontal rows; 18 or 20 
(rarely 16) scales around the caudal peduncle. 

Gill rakers in the outer series on the first ceratobranchial 
9-11, modes 10 and 11. 

Outer series of teeth in both jaws composed mainly of 
unequally bicuspids, but with a few unicuspids or weakly 
shouldered bicuspids present, in fishes < 80 mm S.L.; how- 
ever, in some specimens of Chetia flaviventris, unicuspids 
predominate in fishes in the upper part of that size-range. In 
specimens >100 mm S.L. the outer teeth are predominantly 
unicuspid, with a few very weakly shouldered bicuspids also 
present. Inner tooth rows of both jaws arranged in a double 
series anteriorly and anterolateral^, reducing to a single row 
laterally and posteriorly. In one species (Chetia gracilis) at 
least some specimens have the two anterior median teeth in 
the outer row of inner teeth enlarged and displaced anteriorly 
relative to the other teeth in that row. 

Pre-shank length of the maxilla equal to its shank length 
(see p. 34). Height of the premaxillary alveolar process 
73-77% of the height of the entire ascending process (see 
p. 35). 

Except in one species, the lower pharyngeal bone is not 
enlarged, and the median tooth rows are composed of 
bicuspid teeth only a little coarser than their lateral conge- 
ners. In the exceptional species, C. mola (see Balon & 
Stewart, 1983; fig. 12) the bone is greatly hypertrophied and 
massive, with all but a few of its laterally situated teeth 
enlarged and molariform or submolariform. 
Anal fin spots usually small and fairly numerous (7-15) but in 
one species, C. brevis, there are only 3 or 4 large spots. 
Chetia 



Acknowledgements. To Paul Skelton go my sincere thanks for the 
many discussions we have had on the subject of serranochromine 
cichlids, and for providing me with excellently documented material 
and field data. Once again it is a pleasure to thank Mike Bruton, 
Director of the J.L.B. Smith Institute for his hospitality and the fine 
working facilities provided by the Institute, Huibre Tomlinson for her 
tenacity and good humour when producing the typescript, and Elaine 
Heemstra for her excellent art-work. 



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Museum und Instilut 61: 221-272. 
I .ifin. K.F. 1991. Functional morphology, pp. 129-150. In: Keenleyside, M. 

(Ed.) Cichlid Fishes. Chapman & Hall, London. 
Lippitsch, E. 1991. Comparative investigation on scale characters in cichlids. 

Musee Royal de L'Afrique Centrale, Tervuren, Annales Sciences Zoologique 

263: 97-102. 
Oliver, M.K. 1984. Systematics of African cichlid fishes: determination of the 

most primitive taxon, and studies of the haplochromines of Lake Malawi 

(Teleostei, Cichlidae). Ph.D. Thesis, Yale University. 
Poll, M. 1967. Contribution a la faune ichthyologique de 1'Angola. Museu do 

Dundo, Publicacoes Culturais No. 75: 1-381. 

1986. Classification des Cichlidae du lac Tanganika. Tribus, genres et 

especes. Academie Royale de Belgique; Memoires de la Classe des Sciences, 
collection in 8°, 2e series, 45 (2): 1-163. 

Regan, C.T. 1920. The classification of the fishes of the family Cichlidae. I The 
Tanganyika genera. Annals and Magazine of Natural History (9)8: 632-639. 

1922. The classification of the fishes of the family. Cichlidae. II. On 

African and Syrian genera not restricted to the Great Lakes. Annals and 
Magazine of Natural History (9)10: 249-264. 

Skelton, P.H. 1993. A complete Guide to the Freshwater Fishes of Southern 
Africa. Southern Book Publishers. Halfway House, South Africa. (In press). 

Stiassnv, M.L.J. 1989. A taxonomic revision of the African genus Tylochromis 
(Labroidei, Cichlidae); with notes on the anatomy and relationships of the 
group. Musee Royal de I'Afrique Centrale, Tervuren; Annales Sciences 
Zoologiques 258: 1-161. 

1990. Tylochromis, relationships and the phylogenetic status of the 

African Cichlidae. American Museum Novitates no. 2993: 1-14. 

Trewavas, E. 1961. A new cichlid fish in the Limpopo basin. Annals of the 
South African Museum 46(5): 53-56. 



44 P.H. GREENWOOD 

1964. A revision of the genus Serranochromis Regan (Pisces, Cichlidae). 1963. Zur Klassifikation de Cichlidae, am Beispiel der Gattungen 

Musee Royal de I'Afrique Centrale, Tervuren; Annates, serie in -8°, Sciences Tropheus, Petrochromis, Haplochromis und Hemihaplochromis n.gen. 

Zoologique no. 125: 1-58. (Pisces, Perciformes). Senckenbergiana Biologica 44(2): 83-96. 
1983. Tilapiine fishes of the genera Sarotherodon, Oreochromis and Winemuller, K.O. & Kelso-Winemiller, L.C. 1991. Serranochromis alius , a new 

Danakilia viii + 583p. British Museum (Natural History), London. species of piscivorous cichlid (Teleostei:Perciformes) from the upper Zam- 

Wickler, W. 1962. Ei-Attrappen und Maulbriiten bei Afrikanischen Cichliden. bezi river. Copeia 3: 675-686. 

Zeitschrift fur Tierpsychologie 19(2): 129-164. 






Bull. not. Hist. Mus. (Zool.) 59(1): 45-81 Issued 24 June 1993 

A revision of Danielssenia Boeck and Psammis 
Sars with the establishment of two new genera 
Archisenia and Bathy psammis (Harpacticoida: 
Paranannopidae) 

RONY HUYS* 

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5 BD 

J. MICHAEL GEE 

Plymouth Marine Laboratory, Prospect Place, The Hoe, Plymouth PL1 3DH 

CONTENTS 



Introduction 46 

Materials and Methods 46 

Systematics 47 

Family Paranannopidae Por, 1984 47 

Genus Archisenia gen . nov 47 

Archisenia sibirica (Sars, 1898) comb, nov 47 

Remarks 51 

(i) Synonymy 51 

(ii) Autapomorphies 60 

(iii) Intersexuality 60 

Genus Danielssenia Boeck , 1872 62 

(i) Danielssenia fusiformis (Brady , 1880) nee Sars (1910) 62 

(ii) Danielssenia robusta Sars, 1921 62 

(iii) Danielssenia similis Chislenko, 197 1 64 

(iv) Danielssenia typica Boeck, 1872 64 

( v) Amended diagnosis 67 

Genus Bathypsammis gen. nov 67 

Bathypsammis longifurca (Bodin, 1968) comb, nov 69 

Genus Psammis Sars, 1910 72 

(i) Psammis borealis Klie , 1 939 72 

(ii) Psammis kliei : Smirnov, 1946 72 

(iii) Psammis longipes Becker, 1974 72 

(iv) Psammis longisetosa Sars, 1910 72 

(v) Amended diagnosis 75 

Discussion 78 

Key to genera of Paranannopidae 79 

Acknowledgements 80 

References 80 

Synopsis. Archisenia gen. nov. is proposed to accommodate the sibirica-gvoup of the genus Danielssenia Boeck, 
1872. Re-examination of Alaskan material of D. stefanssoni Willey, 1920 has shown the latter species to be a junior 
synonym of D. sibirica Sars, 1898, the type and only species of the new genus. 

Danielssenia robusta Sars, 1921 and Fladenia intermedia (Wells, 1965) are synonymous and consequently F. 
robusta comb. nov. becomes the type species of the genus Fladenia Gee & Huys, 1990. Danielssenia similis 
Chislenko, 1971 is regarded as species inquirenda and the genus Danielssenia is redefined from the type species D. 
typica Boeck, 1872, and two other species (D. quadriseta Gee, 1988 and D. reducta Gee, 1988). 

The status of D. fusiformis (Brady, 1880) nee Sars (1910) is reconsidered and as a result the genus Sentirenia Huys 
& Gee, 1992 is relegated to a junior synonym of Jonesiella Brady, 1880 which is reinstated to accommodate J. 
fusiformis Brady, 1880 and J. eastwardae (Coull, 1971) comb. nov. 

Psammis borealis Klie, 1939 is removed from the genus Psammis Sars, 1910 but retained in the Paranannopidae as 
species incertae sedis. P. longifurca Bodin, 1968 is transferred from Psammis to Bathypsammis gen. nov. The genus 
Psammis is redefined on the basis of the type species P. longisetosa Sars, 1910, and P. longipes Becker, 1974. 

* Visiting Research Fellow of the Institute of Zoology, University of Gent, B-9000 Gent, Belgium. 



46 



R. HUYS AND J. M. GEE 



A detailed redescription of A. sibirica and new illustrations of D. typica, P. longisetosa, P. longipes and B. 
longifurca are provided. 

Intersexuality in copepods and the possible phylogenetic relationships of Danielssenia, Psammis, Fladenia, 
Archisenia gen. nov. and Bathypsammis gen. nov. are briefly discussed. 

A key to the genera of the Paranannopidae is presented. 



INTRODUCTION 



Throughout its taxonomic history up to the late 1980s, the 
genus Danielssenia Boeck, 1872 has served as a repository to 
accommodate different kinds of 'tachidiid' harpacticoid cope- 
pods, in so far that the distinction between this genus and 
Psammis Sars, 1910 almost became no longer tenable (Wells, 
1965, 1967). Gee (1988a) pointed out that differences in 
mandibular gnathobase structure, possibly reflecting different 
diets, could indicate that both genera are trophically isolated, 
but admitted that perhaps more solid morphological evidence 
is necessary to maintain generic distinction. 

The criteria applied by most workers to allocate newly 
discovered species to Danielssenia generally had no phyloge- 
netic significance as they were mainly based on plesiomorphic 
character states (i.e. PI not modified) which are diagnostic of 
a wider group of families. Virtually no effort has been made 
to correctly assess the sexual dimorphism on the swimming 
legs and very little information on detailed mouthpart struc- 
ture has been documented. Both categories of characters 
have nevertheless proved to hold a high phylogenetic infor- 
mation content that can be used to determine relationships 
within the Danielssenia-Psammis core group of genera (Gee 
& Huys, 1990, 1991; Huys & Gee, 1992, in press). 

The impact of Lang's (1944, 1948) classification of the 
Tachidiidae also caused people to lose sight of the relation- 
ships of this core group with taxa beyond the family bound- 
aries. The fact that his artificial subdivision into three 
subfamilies constrained the development of alternative phy- 
logenetic scenarios for a long time is illustrated by the 
ongoing discovery and description of numerous new species 
of Paranannopus Lang, 1936 (placed in the Cletodidae and 
subsequently in the Paranannopidae) and Danielssenia 



(placed in the Thompsonulinae, Tachidiidae) in the post- 
Langian era without any recognition of the close relationship 
between these two taxa. Huys & Gee (1990) inevitably had to 
break down the concept of the Thompsonulinae before they 
could re-allocate the 'danielsseniid genera' to the Paranan- 
nopidae. This group of genera essentially represents the 
continental shelf lineage of the family with a few species that 
secondarily explored deeper habitats (e.g. Leptotachidia iber- 
ica Becker, 1974). Its affinity to the predominantly deepwater 
group, containing Paranannopus and Cylindronannopus 
Coull, 1973, has recently been supported by the redescription 
of Fladenia Gee & Huys, 1990, a possible 'missing link' 
between both lineages (Gee & Huys, 1990). 

This paper is the final contribution to a revision of the 
genus Danielssenia, including the allocation of the sibirica- 
group to a new genus Archisenia, thus reducing the number 
of species previously referred to the genus from 14 to four 
(Table 1). It also presents a revision of the other major genus 
Psammis, resulting in the proposal of a new genus Bathyp- 
sammis. With the revision of these taxa the establishment of 
novel genera draws to a close and, accordingly, a key to 
genera of the Paranannopidae is presented. 



MATERIALS AND METHODS 

Before dissection, the habitus was drawn and body length 
measurements were made from whole specimens temporarily 
mounted in lactophenol. Specimens were then dissected in 
lactic acid, the parts mounted in lactophenol and the prepara- 
tions sealed with glyceel® (BDH Chemicals Ltd, Poole, 
England). All drawings of the specimens were prepared using 
a camera lucida on a Leitz Dialux 20 or Leitz Diaplan 



Table 1 Re-allocation of species previously referred to Danielssenia Boeck, 1872. 



Species previously referred 
to Danielssenia 



Current status 



Reference 



typica Boeck, 1872 
fusiformis sensu (Sars, 1910) 
quadriseta Gee, 1988 
reducta Gee, 1988 
similis Chislenko, 1978 
sibirica Sars, 1898 
stefanssoni Willey, 1920 
fusiformis Brady, 1880 
perezi Monard, 1935 
paraperezi Soyer, 1970 
eastwardae Coull, 1971 
robusta Sars, 1921 
intermedia Wells, 1965 
spinipes Wells, 1967 
minuta Coull, 1969 



Danielssenia typica Boeck, 1872 
Danielssenia typica Boeck, 1872 
Danielssenia quadriseta Gee, 1988 
Danielssenia reducta Gee, 1988 
Danielssenia similis Chislenko, 1978 [sp. inq.] 
Archisenia sibirica (Sars, 1898) comb. nov. 
Archisenia sibirica (Sars, 1898) comb. nov. 
Jonesiella fusiformis Brady , 1880 
Jonesiella fusiformis Brady , 1880 
Jonesiella fusiformis Brady , 1880 
Jonesiella eastwardae (Coull, 1971) comb. nov. 
Fladenia robusta (Sars, 1921) comb. nov. 
Fladenia robusta (Sars, 1921) comb. nov. 
Afrosenia spinipes (Wells, 1967) 
Sentiropsis minuta (Coull, 1969) 



Gee (1988) 

Gee (1988), present account 

Gee (1988) 

Gee (1988) 

present account 

present account 

present account 

present account 

present account 

Huys & Gee (1992), present account 

Huys & Gee (1992), present account 

present account 

Gee & Huys (1988), present account 

Huys & Gee (in press) 

Huys & Gee (in press) 



REVISION OF DANIELSSENIA AND PSAMMIS 



47 



differential interference contrast microscope. The terminol- 
ogy for body and appendage morphology is according to 
Huys and Boxshall (1991). Abbreviations used in the text and 
figures are P1-P6 for thoracopods 1-6; exp(enp)-l (-2,-3) to 
denote the proximal (middle, distal) segment of a ramus. 
Body length was measured from the base of the rostrum to 
the posterior margin of the anal somite. 



SYSTEMATICS 



Family Paranannopidae Por, 1984 

Genus Archisenia gen. nov. 

Synonymy. Danielssenia Boeck, 1872 (part). 

Diagnosis. Paranannopidae. Body large, slightly fusiform 
and dorso-ventrally flattened. Rostrum not hyaline, with 2 
pairs of small sensillae. Somatic hyaline frills minutely den- 
tate. Female genital double-somite with lateral and ventral 
sub-cuticular ridge marking original segmentation; genital 
field with minute copulatory pore and sinusoidal copulatory 
duct leading to transverse seminal receptacle partly located 
anterior to genital slit; P6 with 1 outer plumose seta and 2 
minute spiniform elements. Pseudoperculum hyaline with 
dentate margin. Caudal rami slightly divergent and slightly 
longer than broad. Female antennule 6-segmented; aes- 
thetasc on segment 4; distal 2 segments with heavily pectinate 
spines. Antennary exopod 3-segmented with armature for- 
mula [2-1-3]. Mandibular coxa elongate, with blunt teeth on 
gnathobase; basis with 4 setae; endopod 1 -segmented; exo- 
pod 2-segmented. Maxilliped subchelate with 1 large and 1 
small seta on syncoxa; basis with naked seta on palmar 
margin, endopodal claw with 2 accessory setae. PI exopod 
3-segmented, exp-3 with distal outer spine longer than middle 
outer spine; endopod longer than exopod, 2-segmented, 
enp-2 4.5 times longer than broad, inner seta implanted 
medially. P2-P4 intercoxal sclerites with spinules or setules 
on distal margin; rami 3-segmented; exp-1 with inner seta; 
female P2-P3 enp-2 with small apophysis at outer distal 
corner. Armature formula of P1-P4 as follows: 



Exopod 



Endopod 



PI 


0.1.023 


1.121 


P2 


1.1.223 


1.1.221 


P3 


1.1.323 


1.1.321 


P4 


1.1.323 


1.1.221 



Female fifth pair of legs not fused medially; exopod and 
baseoendopod separate, each with 5 setae, inner seta on 
exopod well separated from remaining 4 setae. 

Male with sexual dimorphism on antennule, PI, P2 endo- 
pod, P3 endopod, P5, P6 and in genital segmentation. 
Antennule 9-segmented, subchirocer; segment 6 very swol- 
len, with aesthetasc. PI inner basal spine less strongly devel- 
oped, segments of rami more slender and spinule rows on 
outer margin of endopod much smaller. P2 enp-1 larger, with 
inner seta transformed into a non-articulating process; enp-2 
without inner seta, outer distal corner attenuated into a long 
apophysis reaching far beyond the distal border of enp-3; 



enp-3 with distal outer spine partially fused to segment, much 
shorter and stronger than in female, with spinules reduced to 
coarse teeth, other setae reduced in size. P3 enp-2 with inner 
distal corner slightly attenuated, outer distal corner attenu- 
ated into a hook-shaped apophysis. Fifth pair of legs fused 
medially; baseoendopod and exopod separate with 2 and 5 
setae, respectively. P6 symmetrical, fused to somite, with 3 
setae each. 

Type species. As a result of the arguments and analysis put 
forward below we regard D. stefanssoni Willey, 1920 as a 
junior synonym of D. sibirica Sars, 1898 and therefore A. 
sibirica (Sars, 1898) comb. nov. is designated as the type 
species. 

Other species. None. 

Etymology. The generic name is derived from the Greek 
prefix archi, meaning first in time and alludes to the primitive 
position in the family. Gender: feminine. 

Archisenia sibirica (Sars, 1898) comb. nov. 

Synonymy. Danielssenia sibirica Sars, 1898; Danielssenia 
stefanssoni Willey, 1920. 

Material examined. 

— National Museum of Natural History (Smithsonian Institu- 
tion), Washington, D.C.: 8 $$ and 1 cf from Point Barrow, 
Nuwuk Lake, Alaska, U.S.A.; collected by R. Lewis et ai, 
August 1 1960, bottom sample A974; identified as D. stefans- 
soni by M.S. Wilson; 1 °. dissected on 13 slides, 1 d" dissected 
on 7 slides, others preserved in alcohol: reg. no. USNM 
204769. 

— Naturhistoriska Riksmuseet, Stockholm: 1 $ and 1 cf 
from East Greenland, Barclay Bay; collected by Jespersen, 
July 14 1932; identified as D. stefanssoni by K. Lang; 
preserved in alcohol; reg. no. Cop. 31. 

Description of female. Body slightly dorso-ventrally flat- 
tened (as for male, Fig. 9B); length 0.97-1.242 mm (x = 
1.15 mm; n = 7); urosome clearly demarcated from prosome. 
Cephalothorax rounded anteriorly, widest near posterior 
margin. Rostrum (Fig. 2A) not hyaline; tapering anteriorly; 
with 2 pairs of sensillae. Free prosomites each with a dorsal 
row of spinules and some sensillae near posterior margin; 
hyaline frill of prosomites minutely dentate. All urosomites 
(Fig. 1A-B) with lateral row of spinules; first to third 
urosomites with dorsal row of spinules, 2 rows dorsally on 
genital somite; ventral spinule row on posterior border of 
genital double-somite and succeeding urosomites, slightly 
anterior to lateral rows. Genital double-somite (Fig. 1A-B) 
with lateral and ventral subcuticular ridge. Genital field (Fig. 
1C-D) with minute copulatory pore posterior to genital slit; 
copulatory duct sinusoidal (Fig. ID) leading to single, trans- 
versely elongate seminal receptacle located at level of genital 
slit; vestigial P6 with 1 plumose seta and 2 spinules (vestigial 
setae?); paired, blind ending, cuticular invaginations poste- 
rior to genital field (Fig. 1C). Hyaline frill of urosomites 
minutely dentate, that of penultimate somite extended to 
form pseudoperculum (Figs. IB, 8C). Anal somite deeply 
incised. Caudal rami (Figs. IE, 5F, 8C) tapering posteriorly, 
slightly longer than broad, with short spinule row medially on 
lateral margin and a latero-ventral spinule row on distal 
margin which also has a large pore near ventral outer corner 
(Fig. IE); seta I minute (Fig. 5F); setae IV & V well 



48 



R. HUYS ANDJ.M. GEE 




Fig. 1 Archisenia sibirica comb. nov. A, Female urosome (excluding P5-bearing somite), ventral view; B, same, dorsal view; C, female 
genital field, ventral view; D, same lateral view; E, caudal ramus, ventral view. 



REVISION OF DANIELSSENIA AND PSAMMIS 



49 



developed, spinulose in distal portion (Fig. IE); seta VII 
tri-articulate (Fig. 8C). 

Antennule (Fig. 2A-B) 6-segmented; segment 1 with 2 
spinule rows on outer margin and a plumose seta at outer 
distal corner. Segment 2 with 5 pinnate and 1 naked setae on 
outer margin and 2 pinnate and 1 naked setae posteriorly 
directed on dorsal margin. Segment 3 with 2 pinnate and 6 
naked setae at outer distal corner. Segment 4 with 6 naked 
setae and an aesthetasc. Segment 5 with 3 pectinate spines, 3 
naked and 2 pinnate setae. Segment 6 with 1 pectinate spine 
and 7 naked setae. 

Antenna (Fig. 2C-D). Coxa with a row of spinules proxi- 
mally. Allobasis with long spinules at base of abexopodal, 
pinnate seta. Exopod 3-segmented with armature formula 
[2-1-3]; distal segment elongate with subterminal row of 
spinules. Endopod with 2 spinule rows on outer margin; 2 
spines, a geniculate seta and a naked seta subdistally (Fig. 
2C); distal margin with a pectinate spine, 4 geniculate setae, a 
small plumose seta (Fig. 2C) and a very small naked seta (Fig. 
2D). 

Mandible (Fig. 3A-B). Coxa (Fig. 3B) elongate, slender, 
with 2 median rows of spinules; gnathobase with bidentate 
and unidentate teeth and a pinnate seta at inner distal corner. 
Palp biramous. Basis (Fig. 3A) with patch of spinules medi- 
ally and 4 pinnate setae on distal margin. Exopod 
2-segmented; proximal segment with 2 pinnate setae laterally 
and a row of large spinules distally; distal segment with 3 
apical setae. Endopod 1-segmented with 3 lateral and 6 distal 
setae. 

Labrum (Fig. 3C) with numerous spinule rows near median 
distal margin of posterior face. 

Maxillule (Fig. 3D). Praecoxal arthrite with 2 juxtaposed 
setae medially on anterior surface and 9 bidentate or pinnate 
spines and 1 naked seta on distal margin. Coxal endite with 5 
armature elements on distal margin. Basal endite with 2 
subdistal setae and 4 setae on distal margin. Rami 
1-segmented and each with 3 setae. 

Maxilla (Fig. 4B). Syncoxa with spinule row at outer 
proximal corner and with 3 endites each with 1 fused and 2 
articulating pinnate spines. Allobasal endite with a fused 
pinnate claw, a pinnate spine and 2 setae. Endopod 
1-segmented with a pinnate spine and 3 setae. 

Paragnaths (Fig. 4A) well developed; with several rows of 
fine spinules laterally and medially; anterior face with coarse 
teeth. 

Maxilliped (Fig. 4C). Syncoxa with numerous spinule rows, 
1 large subterminal and 1 smaller terminal pinnate seta. Basis 
with row of spinules and a naked seta on palmar margin. 
Endopodal claw as long as basis, spinulose distally and with 2 
accessory setae proximally. 

PI (Fig. 5A). Intercoxal sclerite rectangular with 2 groups 
of setules on each side. Coxa with rows of spinules on 
anterior face and outer margin. Basis with row of spinules on 
inner and distal margin and around base of inner pectinate 
spine (Fig. ID) and outer pinnate seta. Exopod 3-segmented, 
| each with row of spinules on outer margin, outer spines 
pectinate, distal outer spine on exp-3 longer than middle 
outer spine. Endopod longer than exopod, 2-segmented; 
proximal segment slightly longer than broad, distal segment 
about 4.5 times longer than broad, inner seta implanted 
medially. 

P2-P4 (Figs. 6A, 7A, 8A). Intercoxal sclerite with row of 
spinules or setules on each side. Both rami 3-segmented, 
equal in length in P2 but with endopod shorter than exopod in 



P3 and P4; all segments with rows of spinules on outer 
margin; P2 and P3 with a large spinule at base of each inner 
seta on enp-2 and -3. Exp-1 with inner seta; enp-2 with outer 
distal margin somewhat attenuated. Armature formula of 
swimming legs as in generic diagnosis. 

Fifth pair of legs (Fig. 11D) not fused medially; exopod and 
baseoendopod separate. Baseoendopod with short row of 
spinules at base of exopod and setophore of outer seta; 
endopodal lobe well developed, tapering distally, with 5 
pinnate setae, second outer seta longest. Exopod wider than 
long, boundary with baseoendopod straight, not reaching to 
distal margin of endopodal lobe; with 5 pinnate setae, 4 
grouped together on distal outer margin and 1 well separated 
near inner distal corner. 

Description of male. As in female except for following 
characters. 

Body (Fig. 9). Length 1.008 mm (n = 1); second and third 
urosomites not fused and ornamental spinules on urosome 
somewhat more robust (Fig. 11 A). 

Antennule (Fig. 10) 9-segmented, subchirocer with 6th 
segment very swollen, geniculation between 6th and 7th 
segments. Segmental fusion pattern: I, II, III— VIII, IX-XII, 
XIII, XIV-XX, XXI-XXIII, XXIV-XXV, XXVI-XXVIII. 
Armature formula: [1, 1, 11,8, 1, 14+ae, 4, 3, 8]. Segment 6 
very swollen with a complicated pattern of ridges and teeth 
on anterior surface (Fig. 10C-D). Segment 7 with 4 setae, 3 
of which sagittiform, on anterior surface (Fig. 10E). 

PI. Coxa with fewer spinule rows on anterior surface. 
Inner spine on basis without spinule row at base; inner spine 
less well developed and with finer spinules (Fig, 5E) than in 
female (Fig. 5D). Segments of both rami (Fig. 5B) more 
elongate than in female. Spinules on outer and distal margin 
of endopod segments much finer than in female, particularly 
on distal margin of enp-1 (Fig. 5C). 

P2 (Fig. 6B-C). Basal pedestal and articulating surface of 
endopod enlarged. Enp-1 much larger than in female and 
inner seta transformed into a non-articulating process with a 
flagellate tip; outer spinules small. Enp-2 without inner seta 
or spinule row on outer margin; outer distal corner attenu- 
ated into an apophysis reaching well beyound the distal 
margin of enp-3. Enp-3 (Fig. 6C) reduced in size with no 
outer spinule row; outer distal spine shorter but stouter than 
in female with spinules reduced to coarse blunt teeth; termi- 
nal and inner setae also reduced in size compared to female. 

P3 endopod (Fig. 7B-C). Enp-2 without outer spinule row; 
outer and inner distal corners much more attenuated than in 
female, apophysis at outer corner with hooked tip (Fig. 7C); 
inner seta much smaller than in female. 

P5 (Fig. 11B). Baseoendopods of each leg fused medially; 
not fused to exopod. Endopodal lobe reduced with 2 pinnate 
setae of very unequal length. Exopod with 5 pinnate setae, 
inner seta small, middle seta longest. 

P6 a single plate fused to somite proximally (Fig. 11 A), 
with 3 pinnate setae on each side (Fig. 11C). 



REMARKS 



(i) Synonymy 

The Alaskan material on which the above redescription is 
based, was first described in detail in an excellent paper by 



50 




Fig. 2 Archbenia sibirica comb. nov. A, Rostrum and female antennule (armature omitted); B, female antennule (disarticulated); C, 
antenna, anterior view; D, antennary endopod, posterior view of distal margin. 



REVISION OF DANIELSSENIA AND PSAMMIS 



51 




i"ig. 3 Archisenia sibirica comb. nov. A, Mandibular palp; B, mandibular gnathobase; C, labrum; D, maxillule. 



52 



R. HUYS AND J.M. GEE 




Fig. 4 Archisenia sibirica comb. nov. A, Right paragnath, posterior view; B, maxilla with disarticulated endopod; C, maxilliped. 



REVISION OF DANIELSSENIA AND PSAMMIS 



53 




Fig. 5 Archisenia sibirica comb. nov. A, Female PI, anterior view; B, male PI, protopod and endopod, anterior view; C, male PI, distal 
margin of enp-1 of other side; D, female PI inner basal spine; E, male PI inner basal spine; F, caudal ramus, lateral view. 



54 



R. HUYS ANDJ.M. GEE 




Fig. 6 Archisenia sibirica comb. nov. A, Female P2, anterior view; B, male P2 basis and endopod, anterior view; C, male P2 endopod distal 
segment, posterior view. 




55 



Fig. 7 Archisenia sibirica comb. nov. A, Female P3, anterior view; B, male P3 endopod, anterior view; C, male P3, detail of outer apophysis 
of enp-2. 



56 



R. HUYS AND J.M. GEE 




Fig. 8 Archisenia sibirica comb. nov. A, Female P4, anterior view; B, male P4, contours of endopod; C, pseudoperculum, anal somite and 
left caudal ramus in dorsal view. 



REVISION OF DANIELSSENIA AND PSAMMIS 



57 




Fig. 9 Archisenia sibirica comb. nov. A, Male habitus, dorsal; B, same, lateral. [Sensillae on cephalothorax omitted.] 



58 



R. HUYS ANDJ.M. GEE 




Fig. 10 Archisenia sibirica comb. nov. A, Male antennule (armature omitted); B, male antennule, disarticulated (armature of segment 6 
omitted); C, male antennule segment 6, anterior view; D, same, ventral view; E, male antennule segment 7, anterior view. 



REVISION OF DANIELSSENIA AND PSAMMIS 



59 




Fig. 11 Archisenia sibirica comb. nov. A, Male urosome, ventral view; B, male P5; C, male P6 armature; D, female P5. 



60 



R. HUYS AND J.M. GEE 



Wilson (1966). There are slight discrepancies between these 2 
descriptions which should be noted because of their possible 
phylogenetic significance. The rostrum does articulate with 
the cephalothorax and there are 2 accessory setae at the base 
of the maxillipedal claw. In the female there is no aesthetasc 
on the terminal segment of the antennule and the outer distal 
corner of enp-2 in P2 and P3 is significantly attenuated but 
that of enp-1 is normal (especially compared to the condition 
in Psammis). In the male the antennule is distinctly 
9-segmented with segment 6 being swollen and bearing an 
aesthetasc; the distal outer element of P2 enp-3 is not 
completely fused to the segment but articulates along the 
anterior surface and partially articulates on the posterior 
surface; and there is no sexual dimorphism in P4 enp-2 (Fig. 
8B). 

We have been unable to locate the type, or any other, 
material of D. sibirica and therefore, like Wilson (1966), have 
had to rely on Sars' (1898) original description and figures. A 
study of these has led us to conclude that there are no real 
differences between D. stefanssoni and D. sibirica. We agree 
with Wilson's interpretation of the swimming leg setation. 
The original copy of Sars' paper in our possession also shows 
faint lines where the seta should be on P2-P4 exp-1 and in no 
other species in any paranannopid genus is this seta absent in 
the female when it is present in the male. Although Sars 
(1898) states in his text that the female antennule is 
5-segmented, close scrutiny of his drawing (Plate X, Fig. 4) 
reveals that the terminal segment is at least partially divided 
into two segments and that the arrangement of the pectinate 
spines (3 on segment 5 and 1 at the apex of segment 6) is 
exactly the same as in D. stefanssoni. Further Sars has clearly 
misinterpreted the segmentation of the male antennule in 
that he has combined segments 1 and 2 as shown by the fact 
that his segment 1 bears 2 setae, a condition found in no other 
Paranannopidae. Wilson (1966) concluded that the only real 
difference between D. stefanssoni and D. sibirica was the 
absence in the latter of the distinctive outer distal spine on P2 
enp-3 in the male. However, Sars' drawing of this limb (Plate 
X, Fig. 16) shows only 4 elements on enp-3 instead of 5, as in 
the female, and it is possible that the crucial one (the outer 
spine) is concealed behind the large apophysis on enp-2. This 
interpretation is supported by the fact that, irrespective of the 
degree of modification in males, the number of elements on 
P2 enp-3 is always the same in both sexes, except in the 
genera bearing claviform aesthetascs on the mouthparts (Gee 
& Huys, 1991) where the distal outer spine is further modi- 
fied into a non-articulating apical apophysis. 

These observations, coupled with such factors as similarity 
of size (they are the largest known members of the Paranan- 
nopidae except for Psammis borealis Klie, 1939 whose taxo- 
nomic position is unclear) and the peculiar identical 
distribution of setae on the distal margin of the female P5, 
lead us to the conclusion that the two species are synonymous 
and have a circum-polar distribution. Fig. 13 shows that the 
most easterly record of D. sibirica at Wrangell Island (Yash- 
nov, 1935) is very close to the most westerly record of D. 
stefanssoni on the Chukchi Sea coast of Alaska (Wilson, 
1966). The only record of the species outside the Arctic Circle 
is that of Wells (1965) from Loch Nevis on the west coast of 
Scotland and this must be regarded as doubtful (original 
specimens no longer available for re-examination). Further, 
both D. stefanssoni and D. sibirica have been recorded from 
estuaries and in brackish water, a most unusual habitat for 
members of this family. All other species are found only in 



sublittoral marine habitats although Danielssenia typica 
Boeck, 1872, the other species with a known global distribu- 
tion, has been recorded from the Baltic Sea (Veldre & 
Maemets, 1956; Arlt, 1983), a region of lowered salinity. 

(ii) Autapomorphies of Archisenia 

Turning now to consider the taxonomic status of D. sibirica, 
its distinction from other members of the genus Danielssenia 
was suggested by Lang (1944) who divided Danielssenia into 2 
groups, the typica group and the sibirica group. The latter he 
characterized by: (i) antenna exp-1 with 2 setae; (ii) P4 enp-3 
with 2 inner setae; (iii) the male P2 enp-1 with the inner seta 
transformed into a non-articulating process; (iv) the male P3 
enp-2 with an outer hooked apophysis. This last character is 
now known to occur in all species of Paranannopidae and 
might even be a diagnostic character for a wider group of 
families. The first two characters, although diagnostic of D. 
sibirica in combination, are the plesiomorphic condition in 
the family and are found in a number of other genera. 
Paranannopus, Psammis, Micropsammis Mielke, and 
Bathypsammis gen. nov. also retain 2 setae on exp-1 of the 
antenna (all other genera bear only 1 seta on this segment) 
and Sentirenia Huys & Gee (= Jonesiella Brady, see below) 
and the male of Fladenia retain 2 inner setae on P4 enp-3. 
However, the transformation of the inner seta into a non- 
articulating spine on P2 enp-1 in the male is unique to this 
genus, as are the following autapomorphies: (i) the outer 
extension on P3 enp-2 in both sexes; (ii) the sigmoid, heavily 
sclerotized female copulatory duct; (iii) the sexual dimor- 
phism of the inner basal spine of the male PI. Another 
character of phylogenetic significance is the loss of the inner 
seta on the male P2 enp-2. This character is also found in 
Afrosenia spinipes (Wells, 1967) and is regarded here as a 
product of convergence. A further character which is unique 
but difficult to quantify is the arrangement of the setation on 
the exopod of P5, with the inner seta well separated from the 
remaining setae. It is on the basis of all these characters that 
we have removed D. sibirica to a new genus leaving the typica 
group as the only species group in the genus Danielssenia. 

(iii) Intersexuality 

The male from East Greenland (Fig. 12), collected by Jes- 
persen, proved upon examination to be an aberrant intersex- 
ual specimen. It has the male body facies, including a 
6-segmented urosome (Fig. 12A), a well developed testis and 
vas deferens (however, a spermatophore has not been 
observed), and the male form of the P5 and P6. The 
endopods of P2 and P3 are also modified but differ from the 
typical male condition by the retention of certain female 
features. 

The antennules resemble the female condition in all 
aspects: they are 6-segmented, lack any trace of a genicula- 
tion mechanism and possess the female armature pattern. 
The PI (Fig. 12B) basis and endopod show the same spinule 
arrangement as in the female but size and shape of certain 
spinule rows approach the male condition. The P2 endopods 
are not identical on both sides (Figs. 12C-D) and show a 
combination of male and female characteristics. The proximal 
segment and its basal pedestal are moderately enlarged but 
the spinule at the base resembles the female condition and 
the inner seta is — though being shorter than in the female — 
not transformed into a spinous process. The outer apophysis 



REVISION OF DANIELSSENIA AND PSAMMIS 



61 




ig. 12 Archisenia sibirica comb. nov. Intersex specimen from Greenland. A, Habitus, dorsal view; B, PI, endopodal segments; C, P2, 
endopod of right side; D, P2, middle endopod segment of left side; E, P3 endopod; F, P5. 



62 



R. HUYS AND J. M. GEE 



of the middle segment is distinctly shorter than in the typical 
male and its outer margin might bear spinules as in the 
female; the inner seta — completely missing in the male — is 
represented by a vestigial spine which is either entirely (Fig. 
12C) or partly (Fig. 12D) invaginated. The distal segment is 
almost identical to the female condition. The P3 endopod is 
modified as in the male except that the inner seta of the 
middle segment is distinctly longer than in the typical male 
(but shorter than in the female). The P4 endopod grossly 
resembles the male condition. The P5 also has the basic male 
outline but the endopodal lobe is slightly more pronounced 
and the inner exopodal seta is distinctly longer. 

Intersexuality within the Harpacticoida appears to be very 
rare. Klie (1944) describes a female specimen of Amphias- 
coides debilis (Giesbrecht, 1881) from Helgoland which dis- 
played the male condition for the antennules (i.e. haplocer) 
and the first thoracopods (i.e. modified basis) and the female 
condition for the genital somite and the remaining append- 
ages except for the P2 endopod which combined both male 
and female features. Recently, Moore & Stevenson (1991) 
found that 90% of a population of Paramphiascella hyper- 
borea (T. Scott, 1903) in the vicinity of a sewage outfall in the 
Firth of Forth, Scotland, were intersex specimens. In the 
majority of these the prosome (including the antennules and 
swimming legs) exhibited the female condition whilst the 
urosome had the male condition of 6 distinct somites and a 
plate-like P6, although the P5 was more similar to that of the 
female. At the same site, a small number of intersex speci- 
mens of Stenhelia gibba Boeck, 1864 and Halectinosoma 
similidistinctum Lang, 1965 were also found. Intersexuality is 
more common in other orders of copepods, particularly the 
calanoids Eudiaptomus vulgaris (Schmeil, 1898), Arctodiap- 
tomus (Rhabdodiaptomus) alpinus (Imhof, 1885), Eudiapto- 
mus gracilis (Sars, 1863), Pseudocalanus elongatus (Boeck, 
1864), Calanus hyperboreus Kr0yer, 1838, Paracalanus par- 
vus (Claus, 1863) (Bremer, 1914; Pirocchi, 1940; Cattley, 
1949; Francois, 1949; Conover, 1965; Ianora etal, 1987) and 
cyclopoids Megacy clops gigas (Claus, 1857) and Megacy clops 
viridis (Jurine, 1820) (Mrazek, 1913; Coker, 1938). 

In natural populations the frequency of occurrence of 
intersexuality appears to be very low and may be a result of 
infrequent chromosomal aberrations during embryonic devel- 
opment. In cases of higher incidence, various causes of 
intersexuality have been postulated. Coker (1938) attributed 
it to low temperature during naupliar development; Cattley 
(1949) to parasitism of the developmental stages by the 
marine ectoparasitic dinoflagellate Blastodinium contortum 
hyalinum Chatton; and Moore & Stevenson (1991) argued 
that the very high incidence of intersexuality in the vicinity of 
a sewage outfall strongly implicated some form of chemical 
pollution as the causative factor. 

Genus Danielssenia Boeck, 1872 

Since the publication of Lang's (1948) monograph a number 
of new species have been assigned to the genus Danielssenia 
but recent analyses have shown this to be a heterogeneous 
assemblage. In previous papers (see also Table 1) we have 
removed D. intermedia Wells, 1965 to the genus Fladenia; D. 
perezi Monard, 1935 (syn. D. paraperezi Soyer, 1970) and D. 
eastwardae Coull, 1971 to the genus Sentirenia and propose to 
remove D. spinipes Wells, 1967 and D. minuta Coull, 1969 to 
two other new genera (Gee & Huys, 1990; Huys & Gee, 
1992, in press). This has restricted the genus Danielssenia to 



the following species: D. typica; D. quadriseta Gee, 1988; D. 
reducta Gee, 1988; D. robusta Sars, 1921 and D. similis 
Chislenko, 1971. The status of D. fusiformis (Brady, 1880), 
previously been synonymized with D. typica (cfr. Shen & Bai, 
1956; Gee, 1988b) is reconsidered here. 

(i) Danielssenia fusiformis (Brady, 1880) nee Sars (1910) 

Brady (1880) created the genus Jonesiella to accommodate 
two new species, /. fusiformis (Brady & Robertson) and J. 
spinulosa (Brady & Robertson), and provided illustrated 
descriptions for these species. Brady remarked that both 
species had already been listed in an earlier report (Brady & 
Robertson, 1876) as Zosime (?) fusiformis and Z. spinulosa, 
respectively, and therefore unjustly concluded that both 
authors had to be credited with the authorship. This state of 
affairs has perpetuated in the nomenclature, even to the 
present (e.g. Gee, 1988b), though it is clear that Brady & 
Robertson's species names are mere nomina nuda and only 
Brady (1880) should be cited as the author of both Jonesiella 
species. Norman & Scott (1906) were the first to list J. 
spinulosa Brady, 1880 as a junior synonym of Danielssenia 
typica Boeck, 1872 and also changed J. fusiformis Brady, 
1880 into D. fusiformis. Both species were redescribed and 
illustrated by Sars (1910) who admitted that they were very 
similar. It were also Sars' descriptions that led Shen & Bai 
(1956) to conclude that both species were identical, and after 
careful examination of Sars' material Gee (1988b) formally 
relegated D. fusiformis sensu Sars (1910) to a junior synonym 
of D. typica. There is, however, considerable evidence that 
what Sars (1910) considered to be D. fusiformis in Norway is 
clearly different from Brady's (1880) original material from 
the Scilly Islands. Brady's type material does no longer exist, 
but his illustrations (Plate XL VIII, Figs 1-13) of the female 
antennule, mandible, maxilliped, PI, the fifth legs in both 
sexes and the male endopod P2 leave no doubt that his 
species is identical with D. perezi Monard, 1935, originally 
described from Roscoff and later also recorded from the 
Scilly Islands (Wells, 1968) — the type locality of J. fusi 
formis. Huys & Gee (1992) recently synonymized D. para- 
perezi Soyer, 1970 with D. perezi and established a new genus 
Sentirenia to include the latter species and D. eastwardae 
Coull, 1971. Sentirenia Huys & Gee, 1992, therefore, has to 
be relegated to a junior synonym of Jonesiella, thus encom- 
passing the type species /. fusiformis Brady, 1880 nee Sars 
(1910) (syn. nov.: Danielssenia perezi Monard, 1935; D. 
paraperezi Soyer, 1970) and/, eastwardae Coull, 1971 comb. 
nov. 

Thompson's (1893) illustration of the female antennule 
suggests that his record of/, fusiformis from Liverpool Bay is 
correct. Re-examination of specimens (7 $9 labelled D. 
fusiformis; Norman collection, reg. no. 1911.11.8.43561-565, 
gift from T. Scott; October 1899) collected in the Firth of 
Clyde indicates that the species might be distributed along the 
entire west coast of Britain. Lang's (1936a, b) specimens from 
the Oresund and Spitzbergen clearly belong to D. typica. All 
other records of D. fusiformis have to await confirmation (see 
list in Lang, 1948). 

(ii) Danielssenia robusta Sars, 1921 

Lang (1948) was of the opinion that D. robusta (and D. 
perezi) probably would require the definition of additional 
species groups inside the genus but as the males were still 






REVISION OF DANIELSSENIA AND PSAMMIS 



63 




r ig. 13 Distribution map of Danielssenia sibirica (circles) and D. stefanssoni (stars). Records of 1. Sars (1898); 2. Yashnov (1935); 3. Wells 
(1965); 4. Willey (1920); 5. Jespersen (1939); 6. Wilson (1966). Arctic Circle shown by dashed line. 



64 

unknown at that time he regarded such an allocation as being 
premature. We have re-examined Sars' material of this 
species from RisOr, Norway (13 9 $ and 1 copepodid V stage; 
Zoologisk Museum, Oslo, Reg. No. F20257) and found the 
following significant discrepancies from the original descrip- 
tion of Sars (1921): (i) there is an inner seta on exp-1 of 
P2-P4; (ii) the inner element on enp-1 of P2-P4 is a pinnate 
spine; (iii) the P5 baseoendopod has 4 setae, the second inner 
one being much smaller than the others; (iv) the P5 exopod is 
fused to the baseoendopod on the posterior surface. Further, 
we have made a detailed comparison of these females with 
the recently discovered female of D. intermedia (which was 
assigned to the genus Fladenia by Gee & Huys (1990)) and 
have found them to be identical. Therefore D. robusta must 
be referred to the genus Fladenia whose type species now 
becomes F. robusta (Sars, 1921) comb. nov. as this has 
priority over F. intermedia (Wells, 1965). F. robusta has also 
been recorded from the Mediterranean by Por (1964), who 
found one female in 470 m off the coast of Israel, and Soyer 

(1970) who found 18 adult females at depths ranging from 50 
to 420 m in the vicinity of Banyuls-sur-mer. Both authors 
state in their text that these specimens agree exactly with the 
original description. However, Por's (1964) Fig. 73 does show 
an inner seta on exp-1 of what is probably P4 (this limb is 
labelled PI by Por but cannot possibly be so as the endopod is 
3-segmented). This, taken in conjuction with his figure of the 
P5 (1964, Fig. 74) leaves little doubt that the Mediterranean 
material can be assigned to F. robusta, thus giving this species 
a Boreo-Mediterranean distribution similar to that of Jone- 
siella fusiformis (see Huys & Gee, 1992). 

(iii) Danielssenia similis Chislenko, 1971 

Chislenko (1971) distinguished D. similis from D. typica on 
the basis of the following characters: (i) Size, the specimen 
drawn in his Fig. 1 is approximately 0.9 mm long; (ii) a 
maxilliped with only 1 seta on the syncoxa and a somewhat 
longer seta on the basis; (iii) the sexual dimorphism on P2 
endopod, with the loss of the inner seta on enp-1 and of 1 seta 
on enp-3. The character of size is of no particular significance 
as it is within (but near the upper limit of) the size range of D. 
typica given by Gee (1988b). Similarly, the absence of a large 
seta on the basis of the maxilliped is of doubtful significance 
as this seta can be easily dislodged during dissection, as was 
the case in Sars' (1910) description of D. typica (see Gee, 
1988b). The differences in sexual dimorphism of the male P2 
endopod are more difficult to assess from drawings alone. 
However, it is highly improbable that the inner seta on P2 
enp-1 is missing in the male when it is present in the female as 
this condition is found in no other member of this genus or 
indeed of the family as a whole. The same goes for a 
reduction in the number of setae on enp-3. In Danielssenia, 
the 2 terminal setae on this segment are very reduced and 
implanted close together and it is conceivable that Chislenko 

(1971) has combined these 2 fine setae and drawn them as 
one broad one. We believe that D. similis is referable to D. 
typica which has been shown to be the most variable species 
in the genus (Gee, 1988b) but without being able to examine 
topotype material we must regard it as a species inquirenda. 

(iv) Danielssenia typica Boeck, 1872 

The following material of the Norman collection (The Natu- 
ral History Museum) has been examined (species name given 



R. HUYS AND J. M. GEE 
on the original museum label presented in parentheses): 

1911.11.8.43451-470: vial containing > 400 specimens, 
mostly 99> a gift from T. Scott; collected near Duke Buoy, 
Plymouth, 01 August 1889; 

1911.11.8.43471-490: vial containing 23 99 and 1 Q\ a gift 
from T. Scott; collected from Varanger Fjord, East Finmark, 
Norway, 1890; 

1911.11.8.43491-510: vial containing 31 99 and 3 cfd\ a 
gift from T. Scott; collected from Vadso, East Finmark, 
Norway, 03 July 1890; 

1911.11.8.43511-530: vial containing 39 specimens (D. 
typica), a gift from T. Scott; collected in Trondhjem Fjord, 
Norway, 1893; 32 9$ belong to D. typica, the other 7 $$ 
belong to two different species of Halectinosoma; 

1911.11.8.43531-540: vial containing 16 specimens (D. 
typica), a gift from T. Scott; collected from Inchkeith in Firth 
of Forth, October 1895. None of these specimens belongs to 
D. typica, instead the vial contained Bradya sp. (2 99' 8 
copepodids), 2 cfcf Robertsonia tenuis (Brady & Robert- 
son), 1 9 Idomene coronata (T. Scott) and 3 99 OI a 
Fladenia-Uke paranannopid; 

1911.11.8.43541-560: vial containing > 1000 specimens, 
mostly 99- a gift from T. Scott; collected from Karnes Bay, 
Isle of Cumbrae, 1888; a second lot of about 200 specimens 
from the same locality is registrated under no. 1900.3.29.274; 

191 1.11. 8. M. 2299: 1 9 dissected on slide (Jonesiella spinu- 
losa), dried out; collected in Trondhjem Fjord, Norway, 
1893; 

1911. 11. 8. M. 2301: 43 specimens mounted in toto on slide 
(Jonesiella spinulosa), dried out; collected near Duke Buoy, 
Plymouth, 02 August 1889; 

1911. 11. 8. M. 2300: 8 specimens mounted in toto on slide 
{Jonesiella spinulosa), dried out; collected from Vadso, East 
Finmark, Norway, 1890; 

1900.3.6.644: 5 99 mounted in toto and 3 99 (one 
belonging to Halectinosoma sp.) dissected on slide (Jonesiella 
spinulosa); collected in Trondhjem Fjord, Norway, 1893. 

Gee's (1988b) redescription of D. typica is updated here by 
the following observations and illustrations (Figs 14-16) 
based on specimens from Duke Buoy (closest to type local- 
ity): 

Somatic hyaline frills of pedigerous and abdominal somites 
minutely dentate (Fig. 14A) except for the dorsal frill of 
P5-bearing somite which is deeply incised, forming rectangu- 
lar lappets (Fig. 14A, B). Frill of cephalothorax smooth. 
Dorsal transverse spinule rows are found only on thoracic 
somites bearing P3-P5, the genital double-somite and second 
abdominal somite. Genital double-somite with continuous 
transverse chitinous rim dorsally, laterally and ventrally, 
marking original segmentation (Figs. 14A, D; 16D-E). 
Pseudoperculum (Figs. 14E-F) formed by deeply incised 
posterior extension of penultimate somite. Pattern of caudal 
rami setae as in Figs. 14E-F. 

Rostrum (Fig. 161) large, hyaline, with 2 pairs of minute 
sensillae; typically deflected (Figs. 14A-C). 

Male antennule (Fig. 15G) 8-segmented or indistinctly 
9-segmented; distal 2 segments very small and largely fused. 

Mandible with blunt teeth and a single pinnate seta on 
gnathobase (Fig. 15 A). Palp with short, equally long, 
1-segmented rami (Fig. 15B); basis with row of very long 
setules proximally, inner margin with 1 short and 2 long 
setae; endopod with 2 lateral and 6 apical setae; exopod with 




65 



ig. 14 Danielssenia typica. Female: A, habitus, dorsal; B, rostrum and anterior part of cephalothorax, ventral; C, same, lateral; 

D, pleurotergite of P4-bearing somite, P5-bearing somite with fifth thoracopod and genital double-somite, lateral; E, pseudopercuk 
anal somite and left caudal ramus, lateral; F, same, dorsal. [Incised hyaline frill of P5-bearing somite arrowed in A and D.l 



66 




Fig. 15 Danielssenia typica. A, Mandible, gnathobase; B, mandible, palp; C, maxillula, posterior, showing disarticulated praecoxa, coxa and 
palp; D, maxilla, showing disarticulated syncoxa, basis and endopod; E, maxilliped, anterior; F, maxilliped, posterior; G, male antennule 
(armature ommited). [Tubular setae arrowed in C-D.] 



REVISION OF DANIELSSENIA AND PSAMMIS 



67 



row of long setules at 1/3 distance from the proximal margin, 
and 1 lateral, 1 subapical and 2 apical setae. 

Maxillule (Fig. 15C). Praecoxal arthrite with 9 spines and 1 
tubular seta around the distal margin, and 2 geniculate 
tubular setae on the anterior surface; coxal endite with 1 
pinnate spine, 1 setulose claw, 1 smooth setae and 3 tubular 
setae; basal endites closely set, proximal with 2 setae, distal 
with 2 setae and 2 spines; rami with 3 setae each. 

Maxilla (Fig. 15D). Praecoxal endite with 2 unilaterally 
pinnate spines and 1 basally fused spine bearing tubular 
extension. Coxal endites with 1 spine and 2 tubular setae 
each. Allobasis with 2 articulating claws, 1 pinnate spine and 
1 tubular seta. Endopod with 1 tubular seta, 1 spine and 2 
pinnate setae. 

Maxilliped (Figs 15E-F). Syncoxa with 1 large setulose 
spine posteriorly and 1 smaller pinnate seta anteriorly. Basis 
with anterior row of long spinules and small, pinnate seta on 
palmar margin. Endopodal claw with 2 accessory setae. 

Intercoxal sclerites of P2-P4 (Fig. 16H) not U-shaped and 
provided with large spinules on anterior surface (as in Archis- 
enia). 

P2 endopod of male (Figs 16A-B). Inner setae of proximal 
and middle segments reduced compared to the female. Outer 
apophysis of middle segment very large, reaching far beyond 
the distal segment. Inner setae of distal segment spiniform 
and stouter than in the female; inner terminal seta reduced, 
with spatulate tip; outer terminal seta represented by small 
setule; outer spine curved at tip and standing on cylindrical 
process. 

P3 endopod of male (Fig. 16C) with acutely recurved 
process anteriorly at outer distal corner of middle segment. 

Genital field with minute copulatory pore (Fig. 16D) 
leading via short sclerotized duct to multi-chambered seminal 
receptacle (Figs 16F-G). Copulatory duct entering unpaired 
ventral chamber leading dorsally to paired reservoirs both 
anteriorly and posteriorly. Anterior reservoirs largest and 
extending to posterior part of P5-bearing somite (Fig. 16D). 
P6 in female represented by 1 plumose seta and 2 minute 
spiniform elements (Fig. 16E). P6 of male with 1 plumose and 
1 pinnate seta (Fig. 16J). 

Remark. Shen & Bai (1956) pointed out that either 1 or 2 
setae can be found on the middle endopod segment of P2, 
however, their figured specimen with 2 setae on this segment 
(Plate XI, Fig. 86) is almost certainly an aberrant case. The 
same applies to the armature of the baseoendopod of the 
male specimens reported on by Gee (1988b) where a 'vari- 
able' number of setae can be found; all setation patterns 
diverging from the typical bisetose condition are aberrations 
caused by abnormal copepodid development. 

(v) Amended diagnosis 

As a result of the arguments put forward above, the genus 
Danielssenia now contains only 3 well defined species and we 
have re-diagnosed the genus accordingly: 

Paranannopidae. Body variable in size, slightly fusiform 
and dorso-ventrally flattened. Rostrum hyaline, large, typi- 
cally deflexed, with 2 pairs of small sensillae. Somatic hyaline 
frills minutely dentate except for deeply incised frill on dorsal 
margin of P5-bearing somite. Original segmentation of 
female genital-double somite marked by complete sub- 
cuticular ridge; genital field with small copulatory pore; short 
copulatory duct leading to seminal receptacle with paired, 



anteriorly directed chambers extending to anterior margin of 
genital double-somite; P6 with 1 outer plumose seta and 2 
minute spiniform elements. Pseudoperculum hyaline with 
deeply incised margin. Caudal rami parallel, broader than 
long, seta I minute. Female antennule 4-segmented; aes- 
thetasc on segment 3; terminal segment with strong pinnate 
spines. Antennary exopod 3-segmented with armature for- 
mula [1-1-3]. Mandibular coxa with blunt teeth and 1 seta on 
gnathobase, basis broad with 3 setae on distal margin; 
endopod 1-segmented; exopod 1-segmented, with 1 lateral 
and 3 distal setae. Maxillule with 3 tubular setae, 1 pinnate 
seta and 1 spine on coxal endite; basal endite with 4 setae and 
1 spine. Maxilla with tubular setae on coxal endites, allobasis 
and endopod; praecoxal endite with 3 pinnate spines. Maxil- 
liped subchelate with 1 large and 1 small seta on syncoxa; 
basis with small pinnate seta on palmar margin, endopodal 
claw with 2 accessory setae. PI exopod 3-segmented, exp-3 
with distal outer spine longer than middle outer spine; 
endopod 2-segmented, enp-2 4 times longer than broad, inner 
seta implanted medially. P2-P4 intercoxal sclerites with 
spinules on distal margin; rami 3-segmented; exp-1 without 
inner seta. Armature formula of P1-P4 as follows: 



Exopod 



Endopod 



PI 


0.1.023 


1.121 


P2 


0.1.(1-2)23 


1.1.221 


P3 


0.1.(1-2)23 


1.1.(1-2)21 


P4 


0.1.(2-3)23 


1.1.(0-1)21 



Female fifth pair of legs not fused medially; exopod and 
baseoendopod separate, each with 4 or 5 setae. 

Male with sexual dimorphism on antennule, P2 endopod, 
P3 endopod, P5, P6, and in genital segmentation. Antennule 
8- to 9-segmented, subchirocer; segment 6 very swollen, with 
aesthetasc. P2 enp-2 with inner seta, outer distal corner 
attenuated into a long apophysis reaching far beyond the 
distal border of enp-3; enp-3 with distal outer spine and 2 
terminal setae very reduced, inner setae spiniform and larger 
than in female. P3 enp-2 with outer distal corner attenuated 
into a recurved apophysis. P5 of each side fused medially; 
baseoendopod and exopod separate with 2 and 4 or 5 setae, 
respectively. P6 symmetrical, fused to somite, with 2 setae 
each. 

Type species. D. typica Boeck, 1872 (by monotypy). [syn.: 
D. fusiformis (Brady, 1880) sensu Sars (1910)]. 

Other species. D. quadriseta Gee, 1988; D. reducta Gee, 

1988. 

Species inquirenda. D. similis Chislenko, 1971. 



Key to species 

1. P2-P3 exp-3 with 2 inner setae, P4 enp-3 with 1 inner seta ... 2. 
P2-P3 exp-3 with 1 inner seta, P4 enp-3 without inner seta 
D. reductaGee, 1988. 

2. P3 enp-3 with 2 inner setae, P5 exopod with 5 setae in both sexes 

D. typica Boeck, 1872. 

P3 enp-3 with 1 inner seta, P5 exopod with 4 setae in both 
sexes D. quadriseta Gee, 1988. 



68 



R. HUYS ANDJ.M. GEE 







Fig. 16 Danielssenia typica. Male: A, P2 endopod, anterior; B, distal segment of P2 endopod, posterior; C, P3 endopod, middle segment, 
anterior; H, intercoxal sclerite P3; J, sixth leg. Female: D, genital double-somite, ventral; E, genital slit and copulatory pore; F, seminal 
receptacle, lateral; G, same, ventral; I, rostrum. 



REVISION OF DANIELSSENIA AND PSAMMIS 

Genus Bathypsammis gen. nov. 

Synonymy. Psammis Sars, 1910 (part.). 

Diagnosis. Paranannopidae. Body large, more or less cylin- 
drical. Rostrum not hyaline, with 2 pairs of sensillae, anterior 
pair large. Somatic hyaline frills minutely dentate. Female 
genital double-somite with lateral and ventral sub-cuticular 
ridge, marking original segmentation; copulatory pore 
minute; copulatory duct and seminal receptacle unconfirmed; 
P6 with 2 setae and 1 setule in between. Pseudoperculum 
hyaline, vestigial. Caudal rami divergent, elongate (length 
about 5 times proximal width); with tuft of long setules near 
inner distal corner; dorsal surface with chitinized rim in 
anterior half. Female antennule 4-segmented; aesthetasc on 
segment 3; distal 2 segments with heavily pectinate spines. 
Antennary exopod 3-segmented with armature formula 
[2-1-3]. Mandibular coxa elongate, gnathobase with blunt 
teeth and spinule row; basis broad with 4 setae on distal 
margin; endopod 1-segmented, slightly longer than exopod; 
exopod 1-segmented with 1 lateral and 2 apical setae. Maxil- 
lule without modified spines on coxal endite; basal endite 
with 5 setae. Maxilla without tubular setae; praecoxal endite 
with 3 pinnate spines (1 fused to endite). Maxilliped subche- 
late; armature of syncoxa unconfirmed; basis with naked seta 
on palmar margin, endopodal claw with 2 accessory setae. PI 
with very long outer basal seta reaching to middle of exp-3; 
exopod 3-segmented, exp-3 with distal outer spine longer 
than middle outer spine; endopod shorter than exopod; 
2-segmented, enp-2 as long as enp-1, inner seta implanted at 
1/3 distance from proximal margin. P2-P4 intercoxal sclerites 
with few long setules; rami 3-segmented; exp-1 with inner 
seta; female P2-P3 enp-2 without apophysis at outer distal 
corner. Armature formula of P1-P4 as follows: 



Exopod 



Endopod 



PI 
P2 
P3 

P4 



0.1.023 
1.1.223 
1.1.323 
1.1.323 



1.121 
1.2.121 
1.1.121 
1.1.121 



Female fifth pair of legs not fused medially; exopod and 
baseoendopod fused to form a bilobate plate; exopodal lobe 
with 2 spines and 2 setae; endopodal lobe with 2 setae and 3 
spines, the outer 2 of which are stubby. 
Male unknown. 

Type species. Bathypsammis longifurca (Bodin, 1968) 
comb. nov. 

Other species. None. 

Etymology. The generic name is derived from the Greek 
bathys, meaning deep, and Psammis, probably the most 
closely related genus known in the Paranannopidae. Gender: 
feminine. 

Bathypsammis longifurca (Bodin, 1968) comb. nov. 

Synonymy. Psammis longifurca Bodin, 1968. 

Material examined. From Dr Ph. Bodin: holotype dis- 
sected on 3 slides and now deposited in the collections of The 
Natural History Museum under reg. no. 1992.1091; Bay of 



69 

Biscay, Stn 308 (46°07' N; 05°00' W), depth 3950 m; coll. 
August 13 1963, RJVJob-ha-Zelian. 

Bodin's (1968) excellent original description is supple- 
mented here by the following observations and Figures 
17-18. 

Antennule 4-segmented, third segment homologous to 
segments 3-4 in Archisenia. Distal segment with large, swol- 
len seta anteriorly near proximal corner; as pointed out by 
Bodin this segment is seemingly subdivided by the raised 
insertion site of one of the large pectinate spines (Fig. 18A). 
Armature formula: [1, 8, 14+ae, 16]. 

Mandibular gnathobase (Fig. 17B-C) with 4 long teeth, 
one trifid, slender element and 1 pinnate seta; a comb of 
spinules is present at the base of the smaller teeth. The 
endopod has 1 outer, 1 subapical and 6 apical setae (2 of 
which are fused basally). 

Maxillule (Fig. 17D). Praecoxal arthrite with 9 spines and 1 
tubular seta around the inner margin, and 2 geniculate 
tubular setae on the anterior surface; coxal endite with 4 
setae, pinnate spine and straight spine with defined flexure 
zone and small pore near the apex; basal endite with 5 setae. 

Maxilla (Fig. 18B-C) with praecoxal endite drawn out into 
heavily pectinate spine and bearing 2 articulating elements; 
coxal endites with 1 serrate spine and 2 setae each. Allobasis 
with 1 short and 2 long setae; endopod with 4 setae. 

Maxilliped (Fig. 18D). Syncoxa missing in preparation. 
Basis with naked seta on inner margin, 2 spinular rows on 
anterior surface and another one on posterior surface; 
endopodal claw with 2 accessory setae. 

Intercoxal sclerites of P2-P4 U-shaped (as in Fladenia) and 
provided with few long setules near lateral margins. 

Fifth leg (Fig. 17E) with 2 large tube pores on anterior 
surface. 

Genital field (Fig. 18E) with minute copulatory pore. The 
internal structures of the genital double-somite were 
destroyed during the dissection, so no observations of the 
copulatory duct and the seminal receptacles could be made. 

Pseudoperculum very weakly developed. Distribution of 
caudal rami setae as in Fig. 18F-G; seta III dislodged in both 
rami, insertion site indicated by small socle (Fig. 18G). 

P. longifurca does have certain features in common with P. 
longisetosa and P. longipes, namely: anterior pair of rostral 
sensillae enlarged (Fig. 17A); only 1 lateral seta on both rami 
of the mandible (Fig. 17B); 2 setae on exp-1 of the antenna; 2 
setae on enp-2 of the P2; and fused rami in the female P5. 
However, P. longifurca lacks certain important features 
shared by the other two species, namely: no large strongly 
pinnate seta on the basis of the maxilliped, the seta on this 
segment being small and naked (Fig. 18D); the endopod of 
P2 is not distinctly longer than the exopod; the proximal inner 
seta of P2 enp-2 is not displaced to the posterior surface; the 
inner distal seta of P3-P4 enp-3 is not reduced; and, there is 
no attenuation of the outer distal corner of P2 enp-1. Finally, 
P. longifurca has a number of characters which are not shared 
by the other members of this genus such as: (i) a plume of 
long fine setules at the inner distal corner of the caudal 
ramus; (ii) an outer basal seta on PI which is nearly as long as 
the exopod; (iii) a PI endopod which is shorter than the 
exopod and in which both segments are equal in length; (iv) a 
P5 with peculiar spines on the endopodal lobe and a minute 
outer basal seta; (v) a primitive setal formula for the exopods 
of the swimming legs which is shared only by Archisenia and 
Jonesiella. On the basis of these characters we assign P. 



70 



R. HUYS ANDJ.M. GEE 




Fig. 17 Bathypsammis longifurca comb. nov. Female: A, rostrum; B, mandible; C, mandible, gnathobase; D, maxillule, praecoxa 
disarticulated; E, P5, anterior. 



REVISION OF DANIELSSENIA AND PSAMMIS 




71 



ig. 18 Bathypsammis longifurca comb. nov. Female: A, antennule, distal segment; B, maxilla, allobasis and endopod; C, maxilla, syncoxal 
endites; D, maxilliped, anterior (syncoxa missing); E, genital apertures and copulatory pore (arrowed); F, anal somite and left caudal 
ramus, dorsal; G, caudal ramus, detail of posterior margin. 



72 



R. HUYS AND J.M. GEE 



longifurca to a new genus Bathypsammis which is closely 
related to Psammis. 

Genus Psammis Sars, 1910 

With the removal of P. longifurca to Bathypsammis gen. 
nov., the number of species currently allocated to the genus 
Psammis is reduced to four: P. longisetosa Sars, 1910; P. 
borealis Klie, 1939; P. kliei Smirnov, 1946; and, P. longipes 
Becker, 1974. 

(i) Psammis borealis Klie, 1939 

This species was first briefly diagnosed in 1939 from material 
collected in deep water near Iceland. A more extensive 
description, accompanied by illustrations, was published in 
1941. Any justification for placing this species in Psammis is 
missing from Klie's (1939, 1941) papers, providing instead a 
large number of fundamental differences with the type spe- 
cies P. longisetosa. We have re-examined Klie's type material 
of P. borealis (Cop. 211-215; 4 99, 1 cf, all dissected on 
slides; Zoologisches Museum der Universitat Kiel). The slide 
of the male is somewhat confusing in that there seems to be 3 
mounted antennules which do not show male characteristics 
and only part of one which does have the features of a male. 
Further, the limbs on this slide show no sexual dimorphism 
on either P2 or P3. The genital somite is also missing and the 
only appendage that differs from the slides of the females is 
the P5. The fifth legs of both sexes are exactly as drawn in 
Figs. 4 & 6 in Klie (1941). However, based on the mouthparts 
and the setation of the female thoracopods, and pending 
more information on swimming leg sexual dimorphism, we 
propose to retain this species within the Paranannopidae as 
species incertae sedis. It should be noted here that the 
specimens labelled P. borealis and deposited in the Smithso- 
nian Institution (reg. no. 00231018) by Prof. Dr B.C. Coull 
are not the same genus as that of Klie (1939). This material (2 
99) collected from the North Carolina continental shelf [this 
record is not listed in Coull (1971)] closely resembles 
Pseudotachidius similis T. Scott, 1902 and P. minutus ltd, 
1983. 

(ii) Psammis kliei Smirnov, 1946 

We have been unable to discover the type material of P. kliei 
described by Smirnov (1946) from Henrietta Island (New 
Siberian Islands, East Siberian Sea). However, the recent 
recovery of a specimen from Spitsbergen which we believe is 
referable to this species, indicates that it should be placed in 
another genus close to Psammis and Danielssenia. This will 
be discussed further in a future paper on the Paranannopidae 
of Spitsbergen (Gee & Huys, in prep.). 

(iii) Psammis longipes Becker, 1974 

Material examined. Holotype 9 dissected on 2 slides 
(Becker collection; Zoologisches Museum der Universitat 
Kiel, reg. no. 1009-1010); Peru Trough, R/V Anton Bruun 
Sta. 179, 12°03'S 78°45'W, depth 5000 m, leg. W. Noodt. 

This species is known from the type locality only. The 
following redescription (Figs. 19-20) is confined to structures 
that were misinterpreted or not well illustrated in Becker's 
(1974) original description: 

Mandible (Figs. 19A-B). Gnathobase with multicuspidate, 
elongate teeth descreasing in size dorsally, and with 2 pinnate 



setae near the distal dorsal corner; coxa with large spinules 
around the base of the palp. Basis with 3 setae, middle one 
with shorter spinules. Endopod only slightly longer than 
exopod, with 1 lateral and 3 apical setae; exopod with 1 
lateral and 2 apical setae. 

Maxillule (Fig. 19C-D). Praecoxal arthrite with 9 pinnate 
spines and 1 tubular seta around the distal margin and with 2 
geniculate tubular setae on the anterior surface. Coxal endite 
specialized; armature consisting of 3 tubular setae and 3 
spines; largest (= anterior) spine with broad base, a comb of 
flat spinules along the inner margin and ending in a tubular 
extension; middle spine also swollen at base and with fan of 
non-articulating flat spinules arranged around the apex; pos- 
terior spine with large spinule. Basal endite with 3 plumose 
setae and 1 short spine with tubular extension. Endopod and 
exopod with 3 setae each. 

Maxilla (Fig. 19E). Praecoxal endite with 2 pinnate spines, 
distal one with tubular extension. Coxal endites with 2 spines 
and 1 seta each, distal spine and posterior seta with tubular 
extension. Allobasis with 2 articulating claws and a tubular 
seta on either anterior and posterior surface. Endopod with 1 
simple and 3 tubular setae. 

Maxilliped (Fig. 19F) as described by Becker (1974) except 
that the endopodal claw bears an accessory seta. 

The armature formula given by Becker for the swimming 
legs is erroneous on two points: P3 enp-2 has only 1 inner 
seta, the proximal one shown in his figure being an enlarged 
spinule; P3-P4 exp-3 have and extra element distally, repre- 
senting the reduced inner terminal seta (Fig. 20A-B). 

Fifth leg (Fig. 20C). An incomplete furrow on the posterior 
surface marks the original proximal margin of the endopodal 
lobe. The 3 distal setae of this lobe are multipinnate. 

Genital field (Fig. 20D) with small copulatory pore leading 
via linear duct to bilobate seminal receptacle largely located 
anterior to genital slit. P6 armature represented by pinnate 
seta and 2 minute spinules (vestigial setae?). 

Hyaline frill of all body somites finely dentate; pseudoper- 
culum well developed (Fig. 20E). Pattern of caudal ramus 
setae as in Fig. 20E. 

(iv) Psammis longisetosa Sars, 1910 

Material examined. 

— Zoologisk Museum, Oslo: (a) G.O. Sars collection: 

F20223: 1 9 (in alcohol) and 1 ($ (dissected); collected 

from Farsund (type locality), Norway; 

F20224: vial containing 19 99 and 6 Cfcf; collected from 

RisOr, Norway; 

(b) F20929: 4 99 (2 on slides, 2 in alcohol), 3 cfcf (1 on 

slide, 2 in alcohol); collected by J. A. Berg, deposited by J.M. 

Gee, from Bjornehodebukta (59°42.8'N, 10°32.2'E), 

Oslofjord, 35 m depth, June 1984; 

— The Natural History Museum: 1992.1096: 1 d" (in 
alcohol), 1 9 ( on 6 slides), 1 cf prosome (on 7 slides); 
collected by R. Huys, from Frierfjord-Langesundfjord, 55 m 
depth, spring 1985. 

The original descriptions of P. longisetosa given by Sars 
(1910, 1921) have been supplemented since by a complete 
redescription by Gee (1988a). The single female collected 
from Raunefjorden and figured by Por (1965) in all probabil- 
ity does not belong to P. longisetosa. In addition to the 
differences in the shape and armature of the P5 mentioned by 
Por, substantial discrepancies appear from his illustrations of 
the PI (relative proportions of endopodal segments), last 




73 



g. 19 Psammis longipes. Female: A, mandible, posterior; B, mandible, gnathobase, anterior; C, maxillule, posterior; D, maxitlule, detail 
of coxal endite; E, maxilla, posterior (showing syncoxal spines enlarged); F, maxilliped, posterior. [Tubular setae arrowed in C-E.] 



74 



R. HUYS ANDJ.M. GEE 




^■' 



Fig. 20 Psammis longipes. Female: A, P3 endopod, distal segment; B, P4 endopod, distal segment; C, P5, anterior; D, genital apertures and 
copulatory pore (arrowed); E, posterior abdominal somites and left caudal ramus, dorsal. [Vestigial seta arrowed in A-B]. 



REVISION OF DANIELSSENIA AND PSAMM1S 



75 



abdominal somites (ornamentation) and caudal rami (shape). 

Re-examination of P. longisetosa has revealed a number of 
features that were overlooked or misinterpreted in earlier 
descriptions. In many cases these observations have shown an 
astonishing similarity in the detailed structure of the cephalic 
appendages between P. longipes and the type species. 

The rostrum is not hyaline (Fig. 22A); the anterior pair of 
sensillae is enlarged. In the male the antennule is 
9-segmented (Fig. 22A) and the segmental pattern is homolo- 
gous to that of Archisenia. 

Mandible (Fig. 21A-B). The gnathobase has similar multi- 
cuspidate teeth and 2 pinnate setae. The basis has 4 setae; the 
ornamentation of these setae shows that it is either the 
proximalmost or following seta that is missing in P. longipes. 
Both species have the same armature on the rami. 

Labrum (Fig. 22B) with 1 large, median and a pair of 
smaller secretory pores on the anterior surface, and long 
spinules around the distal margin. 

The detailed structure of the maxillule and maxilla is 
exactly the same as in P. longipes, including the presence and 
position of tubular setae and the modifications of the maxillu- 
lary coxal endite. 

The maxillipedal syncoxa has been invariably described as 
possessing a single, very large, spinulose seta, corresponding 
to the posterior seta in P. longipes; the smaller, setulose, 
anterior seta in this species is further reduced to a minute, 
pinnate spine in P. longisetosa (arrowed in Fig. 21C) and 
approaches the length of the largest ornamental spinules, the 
reason why it had been overlooked in previous descriptions. 

The sexual dimorphism on the P2 endopod includes modi- 
fications of the middle and distal segments (Figs 21D-E). The 
anterior, spinous apophysis on the outer margin of the 
proximal segment is not a sexually dimorphic feature since it 
is also found in female specimens. The middle segment is 
drawn out into a large apophysis not reaching to the end of 
the distal segment and provided with an anterior secretory 
pore near the apex; the inner margin has 2 distally serrate 
setae, the proximal one being slightly displaced to the poste- 
rior surface; these setae are distinctly longer in the female. 
The distal segment possesses 4 articulating armature elements 
corresponding to the 2 inner and 2 terminal setae in the 
female; the outer spine in the female is modified in the male 
and replaced by a short, spinous process distally. 

As in P. longipes, the reduced inner terminal seta of P3-P4 
enp-3, represented by a setule, has been overlooked thus far 
(arrowed in Figs 21F-G). In the male the outer distal corner 
of the P3 middle segment is transformed into an acutely 
recurved process (Fig. 21F); the inner seta on this segment is 
distinctly longer in the female. 

The fifth legs of both sexes are as in Figs 22E and F, 
respectively. 

The original segmentation of the female genital double- 
somite is marked by a transverse chitinous rib dorsally and 
ventrally (Fig. 22D). The seminal receptacle is relatively 
small (Fig. 22C); the P6 is represented by 1 plumose seta and 
1 small spinule in the female; in the male the sixth legs are 
fused and symmetrical, and bear 2 naked setae on either side 
(Fig. 22G). 

(v) Amended diagnosis 

Only P. longisetosa and P. longipes are retained in the genus 
Psammis, which is here redefined. 



Diagnosis. Paranannopidae. Body large, slightly fusiform 
and dorso-ventrally flattened. Rostrum not hyaline, with 2 
pairs of sensillae, anterior one large. Somatic hyaline frills 
minutely dentate. Female genital double-somite with lateral 
and ventral sub-cuticular ridge marking original segmenta- 
tion; genital field with minute copulatory pore and linear duct 
leading to transverse seminal receptacle located anterior to 
genital slit; P6 with 1 plumose seta and 1-2 minute spinulose 
elements. Pseudoperculum hyaline with dentate margin. Cau- 
dal rami divergent and longer than broad, tapering slightly. 
Female antennule 4-segmented; aesthetasc on segment 3; all 
segments with pinnate setae and spines. Antennary exopod 
3-segmented with armature formula [2-1-3]. Mandibular 
coxa elongate, with finely pointed teeth and 2 setae on 
gnathobase; basis broad with 3-4 setae on distal margin; 
endopod 1-segmented, equal in length to exopod, with 
strongly reduced armature; exopod 1-segmented, with 1 
lateral and 2 distal setae. Maxillule with 2 large comb-like 
spines and 3 tubular setae on coxal endite; basal endite with 3 
plumose setae, 1 spine and 1 tubular seta. Maxilla with 
tubular setae on coxal endites, allobasis and endopod; prae- 
coxal endite with 2 pinnate spines. Maxilliped subchelate with 
1 large and 1 small seta (both pinnate) on syncoxa; basis with 
long plumose seta on palmar margin, endopodal claw with 1 
accessory seta. PI exopod 3-segmented, exp-3 with distal 
outer spine longer than middle outer spine; endopod at least 
as long as exopod, 2-segmented, enp-2 longer than enp-1, 
inner seta implanted medially. P2-P4 intercoxal sclerites 
without ornamentation; rami 3-segmented; exp-1 with an 
inner seta. P2 endopod distinctly longer than exopod; enp-1 
with outer distal margin attenuated in both sexes; enp-2 with 
1 inner margin seta and 1 seta implanted on posterior surface. 
Inner distal seta enp-3 P3-P4 extremely reduced and repre- 
sented by setule. Armature formula of P1-P4 as follows: 



Exopod 



Endopod 



PI 


0.1.023 


1.121 


P2 


1.1.123 


1.2.221 


P3 


1.1.223 


1.1.321 


P4 


1.1.223 


1.1.221 



Female fifth pair of legs not fused medially; exopod and 
baseoendopod fused to form a bilobate plate; exopodal lobe 
with 4-5 setae, endopodal lobe with 5 setae. 

Male with sexual dimorphism in antennule, P2 endopod, 
P3 endopod, P5, P6 and in genital segmentation. Antennule 
9-segmented, subchirocer; segment 6 swollen, with aes- 
thetasc. P2 enp-2 with long outer apophysis not reaching to 
distal margin of enp-2; enp-3 with outer spine transformed 
into non-articulating process, distal setae reduced and inner 
setae enlarged compared to the female. P3 enp-2 with outer 
distal corner attenuated into a recurved apophysis. Fifth pair 
of legs not fused medially; endopodal lobe with 2 spines, 
exopod with 4 setae/spines. Sixth legs symmetrical, fused to 
somite, with 2 setae each. 

Type species. P. longisetosa Sars, 1910 (by monotypy). 

Other species. P. longipes Becker, 1974. 

Gee (1988a) concurred with Wells' (1967) opinion that a 
generic distinction between Danielssenia and Psammis on the 
base of P5 segmentation alone can hardly be justified. 



76 



R. HUYS ANDJ.M. GEE 




Fig. 21 Psammis longisetosa. Female: A, mandible, gnathobase; B, mandible, palp; C, maxilliped, anterior (small seta on syncoxa arrowed). 
Male: D, P2 endopod, anterior; E, P2 endopod, middle and distal segments, posterior; F, P3 endopod (small seta arrowed); G, P4 endopod 
(small seta arrowed). 



REVISION OF DANIELSSENIA AND PSA MM IS 



77 




ig. 22 Psammis longisetosa. Male: A, antennule and rostrum (armature omitted); F, P5, anterior; G, P6. Female: B, labrum, anterior; 
C, genital apertures and copulatory pore; D, genital double-somite, ventral; E, P5, anterior. 



78 



R. HUYS AND J. M. GEE 



However, Gee also pointed out that the mandibular gna- 
thobase in all Psammis species bears long, relatively fine, 
sharply pointed and widely separated teeth compared to the 
species of Danielssenia where these teeth are short, stout, 
blunt and closely set. On the base of this difference he 
suggested that both genera probably utilize different food 
items and to a certain extent are trophically isolated. In 
combination with the fused rami in the female P5, this 
evidence was considered as sufficient to maintain Psammis'' 
separate generic status. Close examination of the mouthparts 
in P. longisetosa and P. longipes and comparison with D. 
typica has now revealed several other characters that can be 
used to distinguish both genera. Unique features for Psammis 
are the specialized comb-like spines on the coxal endite of the 
maxillule, the presence of only two spines on the praecoxal 
endite of the maxilla, and the extremely enlarged, spinulose 
seta on the maxillipedal basis. The presence of tubular setae 
and modified spines with tubular extensions on the maxillule 
and maxilla is a character that is shared by both genera 
though the precise number is not identical. It is conceivable 
that these specialized structures might perform a sensory role 
(as chemo- or probably mechanoreceptors) in remote food 
detection and/or manipulation. Both genera are predomi- 
nantly found in the upper flocculent layer of muddy sub- 
strates where selection of food-particles probably requires a 
different mechanism. This could be particularly true for 
deepwater bottoms (fjords, abyss) where either turbidity is 
high or the proportion of suspended food-particles might fall 
below a subsistence level. The unique specialization of the 
mandibles, maxillules and maxillae might be viewed collec- 
tively as the result of a different dietary discrimination 
mechanism based on successful remote selection of food 
particles and thus avoiding the unnecessary high energy costs 
of rejecting unsuitable items upon initial capture. It is noted 
here that the claviform aesthetascs found on the mouthparts 
of certain other Paranannopidae (Gee & Huys, 1991) are not 
homologous to the tubular setae or modified spines bearing 
tubular extensions. 

Another unique apomorphy of Psammis is illustrated by 
the setation pattern on the endopods of P3 and P4 (Fig. 23). 
The ancestral condition of P3 enp-3 is shown by e.g. Archise- 
nia and consists of 1 outer spine (a), 2 distal spines (b-c) and 
3 inner setae (d-f). This full complement of armature ele- 
ments is also found in Psammis but is obscured by modifica- 
tions in the distal part of the segment. The extreme reduction 
of the inner terminal spine (c) and the distad displacement of 
the distal inner seta (d) are the main reasons why the setal 
formula was erroneously cited as 221 (or 121 in P4) in 
previous descriptions. The distal elements expressed in this 
formula are b and d, rather than b and c. The spiniform and 
pinnate nature of seta d in Psammis did certainly contribute 
to this misunderstanding. The reduced condition in Bathy- 
psammis (Fig. 23) has not evolved from the Psammis pattern 
but resulted through the loss of 2 inner setae. It is impossible 
to determine which seta (d, e or f) has been retained in B. 
longifurca. 

Both species of Psammis can be differentiated by the 
number of setae on the mandibular basis (3 in longipes, 4 in 
longisetosa), the length of the anterior seta on the syncoxa 
which is distinctly longer in P. longipes, the ratio of endopod 
length to exopod length in PI to P3 being much higher in P. 
longipes, the number of setae on the § P5 exopod (4 in 



longisetosa, 5 in longipes), and the gross difference in body 
size (± 550 u.m in longisetosa, ± 900 nm in longipes). 



DISCUSSION 

Within the Paranannopidae, aesthetascs on the mouthparts 
are a powerful synapomorphy for separating a number of 
genera which have recently been created or redefined, viz. 
Jonesiella (cf. Huys & Gee, 1992), Paradanielssenia , Microp- 
sammis, Telopsammis and Leptotachidia (cf. Gee & Huys, 
1991), Sentiropsis and Peltisenia (Huys & Gee, in press). The 
absence of such sensory appendages in Archisenia excludes it 
from this lineage and allies it with the more primitive 
danielsseniid genera, namely Fladenia, Danielssenia, Psam- 
mis and Bathypsammis. However, the phylogenetic relation- 
ships amongst these more primitive danielsseniid genera are 
somewhat unclear at the moment particularly with respect to 
the position occupied by Archisenia. The problem is that this 
genus shows a mosaic of primitive plesiomorphic characters 
(6-segmented female antennule; setal formula of legs P2-P4 
with 7.8.8 setae/spines on exp-3 and 5.6.5 setae on enp-3; P5 
with 5 setae on baseoendopod and exopod), but at the same 
time a number of unique autapomorphies in the sexual 
dimorphism on PI basis, P2 enp-1 and P3 enp-2. 

Within this group of genera it is clear that Fladenia is the 
most primitive genus because it retains both vestiges of sexual 
dimorphism involving a difference in the number of elements 
(in this case setae) on the endopod of P3 and P4 (Gee & 
Huys, 1990) and a primitive setal formula particularly in the 
exopods of P3 and P4. It is also clear that Danielssenia, 
Psammis and Bathypsammis are linked by a 4-segmented 
female antennule, a reduced number of setae on P4 enp-3 and 
probably by having only 2 setae on the P6 in the male (though 
the latter character cannot be scored for Bathypsammis since 
the male is unknown). Since it has no vestige of P3 and P4 
setal sexual dimorphism and does not show the apomorphies 
of the Danielssenia lineage, it is likely that Archisenia 
diverged from the main evolutionary line after Fladenia and 
probably before the D anielssenia-gr ouying. 

Within the Danielssenia-Psammis-Bathypsammis lineage, 
Danielssenia is considered the most advanced genus on 
account of the loss of a seta on exp-1 of the antenna, the basis 
of the mandible, exp-1 of P2-P4 and enp-2 of P2. Unique 
apomorphies for this genus are the typically ventrally 
deflected rostrum, the blunt teeth on the mandibular gna- 
thobase, and the dorsal, incised, hyaline frill on the 
P5-bearing somite. Another diagnostic character for Daniels- 
senia is illustrated by the shape of the seminal receptacle. 
Multi-chambered receptacles have been described for a num- 
ber of Paranannopidae such as Leptotachidia, Telopsammis, 
Psammis and Paranannopus (Gee & Huys, 1990, 1990) and 
might well be the ancestral state in this family. However, in 
none of these genera the paired anterior chambers are 
elongate, cylindrical reservoirs extending into the posterior 
part of the P5-bearing somite. 

Analysis of the precise relationships within the Danielsse- 
nia grouping is hampered by the absence of male Bathypsam- 
mis. The specialized tubular structures on the endites of the 
maxillule and maxilla provides a robust synapomorphy to link 
Danielssenia and Psammis. A close relationship is also indi- 
cated by the armature of the female sixth legs bearing one 



REVISION OF DANIELSSENIA AND PSAMMIS 



79 




Archisenia 



Psammis 



Bathypsammis 



Fig. 23 Comparison of armature on distal endopod segment of P3 in Archisenia, Psammis and Bathypsammis. 



plumose seta and 2 inner, minute spiniform elements (com- 
pared to 2 setae and 1 setule in between in Bathypsammis), 
and by a detailed comparison of the distal transformations in 
the male P2 endopod. Potential synapomorphies grouping 
Psammis and Bathypsammis are: (i) rostrum with enlarged 
anterior sensillae; (ii) the mandibular exopod with only 1 
lateral and 2 apical setae; (iii) the fusion of the exopod and 
baseoendopod in the female P5. Some species of Danielsse- 
nia, however, also show a reduction in the setation of the 
mandibular exopod (e.g. D. typica), and the fused P5 in 
Bathypsammis might have been evolved convergently, since, 
in other respects, it is very different from the condition in 
Psammis. The rostral character might also be a product of 
convergence since the enlargement of the anterior pair of 
sensillae has evolved independently in a number of other 
deepwater genera such as Paranannopus and Cylindronanno- 
pus. 

Unique apomorphies for Psammis are: (i) reduction of the 
mandibular endopod (1 lateral, 3 apical setae); (ii) the 
specialized comb-like spines on the maxillulary coxal endite; 
(iii) praecoxal endite of maxilla with only 2 spines; (iv) 
extreme development of the posterior seta on the maxillipe- 
dal basis; (v) elongation of P2 endopod, being longer than the 
exopod; (vi) the apophysis on P2 enp-1 in both sexes; (vii) 
reduction of the inner terminal seta on P3-P4 enp-3. In 
Bathypsammis the unique apomorphies are confined to the 
female as the male is unknown: (i) a very long outer basal 
seta on the basis of PI; (ii) a very long caudal ramus with a 
plume of setules on the inner distal corner; (iii) the form of 
the setae on the endopodal lobe of the female P5. 



KEY TO GENERA OF PARANANNOPIDAE 

Remark. This key also includes Psammis kliei Smirnov, 
1946, which will be placed in a genus by itself in a forthcom- 
ing paper (Gee & Huys, in prep.), and the genus Carolinicola 
Huys & Thistle, provisionally assigned to the Paranannop- 
idae by Huys & Thistle (1989). 

1. P4 endopod 3-segmented 4. 

P4 endopod 2-segmented, 1-segmented or absent 2. 

2. Antennary exopod 1-segmented 

Carolinicola Huys & Thistle, 1989. 

Antennary exopod 3-segmented 3. 

3. Body short, robust; caudal rami setae IV and V long and 
spinulose; P5 well developed, covering entire width of thoracic 

somite Paranannopus Lang, 1936. 

Body slender, cylindrical to vermiform; caudal rami setae IV 
and V short and plumose; P5 a minute plate, located midven- 
trally Cylindronannopus Coull, 1973. 

4. P2-P4 exp-1 without inner seta 5. 

P2-P4 exp-1 with inner seta 7. 

5. Antennules without plumose or pinnate spines/ 

setae Sentiropsis Huys & Gee, 1993. 

Antennules with plumose and/or pinnate spines/setae 6. 

6. Caudal ramus with distinct cluster of long setules at the inner 
distal corner; P2 enp-2 with large apophysis in $ (and presum- 
ably in d" also) Psammis kliei Smirnov, 1946. 

Caudal ramus without such cluster; P2 enp-2 with large apophy- 
sis in d" only Danielssenia Boeck, 1872. 

7. P4 exp-3 with 8 setae/spines 8. 

P4 exp-3 with at most 7 setae/spines 13. 

8. P2 enp-2 with 2 inner setae 9. 



80 



R. HUYS AND J.M. GEE 



P2 enp-2 with 1 inner seta 



10. 



REFERENCES 



9. Caudal rami 5 times as long as maximum width; PI endopod 
shorter than exopod; P5 $ with fused exopod and baseoen- 

dopod Bathypsammis gen. nov. 

Caudal rami broader than long; PI endopod longer than exo- 
pod ; P5 $ with separated exopod and baseoendopod 

Jonesiella Brady, 1880. 

10. Body dorsoventrally flattened; caudal rami setae IV and V 
stubby and spiniform; PI enp-1 1.5 times as long asenp- 

2 Peltisenia Huys & Gee, 1993. 

Body not dorsoventrally flattened; caudal rami setae IV and V 
long and setiform; PI enp-1 at most as long as enp-2 11. 

11. Antennule $ 4-segmented; club-shaped aesthetascs present on 
mandible (endopod), maxillule (basis) and maxilla (endopod); 
P2 enp-2 d" without distinct outer apophysis 

Paradanielssenia Soyer, 1970. 

Antennule 9 6-segmented; no club-shaped aesthetascs on 
mouthparths; P2 enp-2 d* with long outer apophysis 12. 

12. Antennary exopod with 1 seta on proximal segment; P3 exp-3 
with 7 setae/spines; P2 enp-3 with inner distal seta transformed 
into large pinnate spine reaching beyond apophysis of enp- 

2 Afrosenia Huys & Gee, 1993. 

Antennary exopod with 2 setae on proximal segment; P3 exp-3 
with 8 setae/spines; P2 enp-3 d with inner distal seta not 
transformed and shorter than apophysis of enp-2 

Archisenia gen . nov . 

13. P2 enp-2 with 2 inner setae Psammis Sars, 1910. 

P2 enp-1 with 0-1 inner setae 14. 

14. Club-shaped aesthetascs present on mandible (endopod), max- 
illule (basis) and maxilla (endopod); P2 exp-3 with at most 6 

setae/spines 15 . 

No club-shaped aesthetascs present on these appendages; P2 
exp-3 with 7 setae/spines Fladenia Gee & Huys, 1990. 

15. PI enp-2 with 2 terminal setae geniculate; P5 $ baseoendopod 
and exopod indistinguishable, with 5 setae; P2 enp-2 d" without 

apophysis; P6 c? with 2 setae 16. 

PI enp-2 with 1 terminal seta geniculate; P5 9 baseoendopodal 
and exopodal lobes indistinguishable, with 3 and 4 setae, 
respectively; P2 enp-2 cf with small apophysis; P6 d" with 3 
setae Micropsammis Mielke, 1975. 

16. Antennule in both sexes with densely opaque, bulbous append- 
age on distal segment P2-P4 exp-2 without inner seta 

Leptotachidia Becker, 1974. 

Antennule in both sexes without densely opaque, bulbous 

appendage on distal segment P2-P4 exp-2 with inner seta 

Telopsammis Gee & Huys, 1991. 



Acknowledgements. The authors wish to thank Dr. Philippe Bodin 
and the Curators of Crustacea at the Zoologisches Museum der 
Universitat Kiel, the Naturhistoriska Riksmuseet Stockholm, the 
Zoologisk Museum Oslo and the National Museum of Natural 
History (Smithsonian Institution) for the loan of material. Dr Geof- 
frey A. Boxshall is acknowledged for his comments on an earlier 
draft of the ms. For the senior author this is communication No. 551 
of the Centre for Estuarine and Coastal Ecology, Yerseke and for the 
junior author the work forms part of the Community Ecology 
Programme of the Plymouth Marine Laboratory. 



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81 



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Bull. nat. Hist. Mus. (Zool.) 59(1): 83-94 



Issued 24 June 1993 



A new species of Syrticola Willems & Claeys, 
1982 (Copepoda: Harpacticoida) from Japan 
with notes on the type species 

RONY HUYS 

Department of Zoology , The Natural History Museum, Cromwell Road, London SW7 5 BD 

SUSUMU OHTSUKA 

Fisheries Laboratory, Hiroshima University, 1294 Takehara-cho, Takehara, Hiroshima 725, Japan 

CONTENTS 



Introduction 83 

Materials and Methods 83 

Descriptions 84 

Family Cylindropsyllidae Sars, 1909 84 

Subfamily Leptopontiinae Lang, 1948 84 

Syrticola Willems & Claeys, 1982 85 

Syrticola intermedius sp . nov 85 

Syrticola flandricus Willems & Claeys, 1982 92 

Remarks 92 

Discussion 93 

Key to species 93 

Acknowledgements 93 

References 94 



Synopsis. A new species of Syrticola Willems & Claeys, 1982 (Harpacticoida: Cylindropsyllidae) is described from 
Okinawa, Japan. Morphological notes on the type species S. flandricus Willems & Claeys, 1982 and a key to the 
species are given. The inadequately described S. trispinosus A. Scott, 1896 is ranked as species inquirenda. The 
diagnosis of the genus is amended and its position in the Cylindropsyllidae re-assessed. Both sexes of S. intermedins 
sp. nov. were found to be infested by early parthenogenetic female stages of an as yet undescribed genus of 
Tantulocarida. 



INTRODUCTION 



The interstitial harpacticoid fauna of Japan is very poorly 
known, as is that of most east Asian countries. The paucity of 
data on marine interstitial species stands in marked contrast 
with the number of studies on subterranean copepods pro- 
duced by workers like Miura and Takashi Ito. In fact, with 
the possible exception of Microsetella norvegica (Boeck, 
1864) only 11 genuinely interstitial harpacticoids have been 
recorded from marine and brackish water habitats in Japan 
(Table 1) and the majority of these was described by the 
latter author's namesake, the late Tatsunori Ito, whose 
activities were mainly focussed on the fauna from Hokkaido 
in the north and the Bonin Islands in the southeast. The only 
other information on mesopsammic harpacticoids is con- 
tained in the papers of Kikuchi (1970, 1972) and Kikuchi & 
Yokota (1984), reporting on species from Lake Hinuma, a 
brackish lagoon near the central east coast of the Japanese 
mainland. 



In the course of a survey of the sandy bottom copepods off 
Nagannu Island, Okinawa (Ryukyu Archipelago) by one of 
us (S.O.), several interstitial harpacticoids were found to be 
infested with tantulocaridans (Huys et ai, in preparation). 
This paper describes a new species of Syrticola Willems & 
Claeys (Cylindropsyllidae) based on two specimens that were 
parasitized by parthenogenetic females of an as yet unde- 
scribed tantulocarid. 



MATERIALS AND METHODS 

Specimens of Syrticola intermedius sp. nov. were collected by 
dredging of a sandy bottom off Nagannu Island, Okinawa, 
South Japan (26° 14' N, 127° 32' E, depth 46.49 m; leg. S. 
Ohtsuka) on 9 April 1992. The dredge (mouth area: 50 cm 
wide x 15 cm high; mesh size 5 mm) was towed along the 
bottom at a speed of 2 knots by the T/V Toyoshio-maru of the 
Hiroshima University for about 5 minutes. Copepods were 



84 



R. HUYS AND S. OHTSUKA 



Table 1 Interstitial harpacticoid copepods reported from marine localities in Japan. 



Species 



Locality 



Reference 



ECTINOSOMATIDAE 

Microsetella norvegica (Boeck, 1864) 
Arenosetella bidenta Ito, 1972 
Noodtiella sp. 

DARCYTHOMPSONIIDAE 

Leptocaris brevicornis (van Douwe, 1904) 



PARAMESOCHRIDAE 
Paramesochra sp. 

LEPTASTACIDAE 

Cerconeotes japonicus (Ito, 1968) 
Paraleptastacus unisetosus Ito, 1972 

CYLINDROPSYLLIDAE 

Arenopontia ishikariana Ito, 1968 
Arenopontia sakagamii Ito, 1978 
Stenocaris intermedia Ito, 1972 
Psammopsyllus imamurai Kikuchi, 1972 

PARASTENOCARIDIDAE 

Parastenocaris hinumaensis Kikuchi, 1970 



Hokkaido 
Hokkaido 
Hokkaido 



Lake Hinuma 1 



Hokkaido 



Hokkaido 
Hokkaido 



Hokkaido 
Bonin Islands 
Hokkaido 
Lake Hinuma 1 



Lake Hinuma 1 



Ito (1968) 

Ito (1972, 1984) 

Ito (1984) 



Kikuchi & Yokota 
(1984) 



ltd (1984) 



Ito (1968, 1984) 
Ito (1972, 1984) 



Ito (1968, 1984) 
Ito (1978) 
Ito (1972) 
Kikuchi (1972) 



Kikuchi (1970) 



1 Brackish lagoon. 



fixed and preserved in 10% neutralized formalin/sea-water. 
Females of S. flandricus Willems & Claeys, 1982 were 
collected by the senior author in different localities along the 
coast of The Netherlands in the course of the biological 
monitoring programme BIOMON. All specimens have been 
deposited in the collections of The Natural History Museum, 
London. 

Specimens were dissected in lactic acid and the dissected 
parts were placed in lactophenol mounting medium. Prepara- 
tions were sealed with glyceel (Gurr®, BDH Chemicals Ltd, 
Poole, England). All drawings have been prepared using a 
camera lucida on a Leitz Diaplan differential interference 
contrast microscope. The descriptive terminology is adopted 
from Huys & Boxshall (1991). Abbreviations used in the text 
are: P1-P6, first to sixth thoracopod. 



DESCRIPTIONS 



Family Cylindropsyllidae 

Subfamily Leptopontiinae Lang, 1948 

The genus Syrticola was established by Willems & Claeys 
(1982) to accommodate the type species S. flandricus Willems 
& Claeys, 1982 and Tetragoniceps trispinosus A. Scott, 1896. 
Previously, the latter species had been considered 'species 
incertcC in the genus Evansula T. Scott and thus placed in the 
subfamily Cylindropsyllinae (Lang, 1948). The close relation- 
ship between Syrticola and Notopontia Bodiou noted by 
Willems & Claeys (1982) was already hinted at by Bodiou 
(1977) who recognized a certain resemblance between T. 
trispinosus and N. stephaniae Bodiou, 1977, and indirectly 
also by Mielke (1982) who described (?) N. galapagoensis, a 



species provisionally placed in Notopontia but subsequently 
allocated to Syrticola (Bodiou & Colomines, 1986; Willems et 
ai, 1987). However, none of these authors has formally 
assigned either of these genera to any of the subfamilies of 
the Cylindropsyllidae recognized at that time. The only 
attempt was that by Bodiou (1977) who suggested that 
Notopontia is closest to Evansula (Cylindropsyllinae) but to a 
certain extent is also related to Arenopontia Kunz and 
Leptopontia T. Scott (Leptopontiinae). 

Lang (1948) subdivided the family into the Cylindropsylli- 
nae, Leptastacinae and Leptopontiinae and a fourth subfam- 
ily, the Psammopsyllinae, was added by Krishnaswamy 
(1956). Recently, the Leptastacinae has been upgraded to full 
family status (Huys, 1993). The diagnostic sexual dimorphism 
displayed on thoracopods 2 and 3 by all genera of the 
Cylindropsyllinae excludes Notopontia and Syrticola from 
this subfamily since their swimming leg sexual dimorphism is 
only slightly developed (and therefore might well have been 
overlooked in Notopontia for which it has been recorded as 
being completely absent). A detailed comparison with the 
Leptopontiinae, currently encompassing Arenopontia, 
Par arenopontia Bodiou & Colomines and Leptopontia, 
reveals a suite of apomorphic characters supporting a sister- 
group relationship between Leptopontia and the Notopontia- 
Syrticola lineage. These characters include: (i) anal 
operculum drawn out into spinous process(es); (ii) outer 
distal corner of caudal ramus produced into backwardly 
directed spinous process; (iii) first antennulary segment ( 
extremely elongated, much longer than second; (iv) mandib- 
ular gnathobase stylet-like with teeth along one side; (v) 
distal exopod segment PI with 3 armature elements (proximal 
outer spine lost); (vi) middle exopod segment PI without 
outer spine (in Syrticola and Notopontia the middle and distal 
segment are fused or have failed to separate); (vii) apical 
spines of distal exopod segments P3-P4 setiform; (viii) sexual 



NEW SPECIES OF SYRTICOLA FROM JAPAN 



85 



dimorphism endopod P3 involving fusion of distal spine to 
segment; (ix) P5 exopod with 3 elements in both sexes. There 
is little evidence that Arenopontia and Pararenopontia share a 
close relationship with this core group, however pending a 
revision of these genera it is preferable to retain them in the 
Leptopontiinae. 

Syrticola Willems & Claeys, 1982 

Diagnosis (amended). Leptopontiinae. Body cylindrical, 
but not particularly vermiform. Hyaline frill of all body 
somites incised. Antennule 6- or 7-segmented in 9- Maxilla 
with one syncoxal endite. Midventral spinous process ante- 
rior to intercoxal sclerite of PI. PI exopod 2-segmented. 
Distal segment PI endopod with 1 geniculate seta and 1 claw. 
Distal segment P3-P4 exopods with 1 outer spine. P2-P4 
endopods 1-segmented in 9> P3 endopod 1- or 2-segmented 
and sexually dimorphic in cf. P5 with fused baseoendopod 
and exopod in both sexes; endopodal lobe drawn out into 
triangular process with 0-1 seta, exopodal lobe a tubercle with 
3 elements. Genital apertures not fused in 9- Anal opercu- 
lum with a series of small spinous processes or one large 
median spike. Caudal ramus seta III inserted proximal to seta 
V. 

TYPE SPECIES. Syrticola flandricus Willems & Claeys, 1982 

Other species. S. trispinosus (A. Scott, 1896), S. galapa- 
goensis (Mielke, 1982), 5. mediterraneus Willems et al. , 1987, 
5. intermedius sp. nov. 

Syrticola intermedius sp. nov. (Figs. 1—4, 5A-C, 6) 

Material examined. Holotype 9 dissected on 8 slides, 
deposited under reg. no. 1992.1075. Paratype a" dissected on 
6 slides, deposited under reg. no. 1992.1076. Drawings based 
on the paratype are Figs. 2E-F, 4D-F, 5A-C, 6A-G; all 
others were drawn from the holotype 9 • 

Female. Body length measured from tip of rostrum to 
posterior margin of caudal rami 485 um (Figs. 1A-B). Maxi- 
mum width 75 [im measured at rear margin of cephalothorax. 
Integument pitted. Pleural areas of cephalothorax not well 
developed so that appendages are clearly exposed in lateral 
aspect (Fig. IB). Posterior margin of body somites (except 
cephalothorax and anal somite) fringed dorsally and laterally 
with finely incised hyaline frill; this frill also present ventrally 
on genital double-somite and abdominal somites (Figs. IB, 
4A-B). Abdominal somites also with transverse spinular row 
in anterior half which is usually concealed beneath the 
hyaline frill of the preceding somite as shown in Fig. 5A. 

Rostrum triangular, with 2 delicate sensillae (as in male, 
Fig. 6B). 

Genital double-somite (Fig. 4B) about as long as wide; 
original segmentation not marked by any external or internal 
cuticular structure; anterior margin with 2 transverse spinular 
rows. Genital apertures located in anterior quarter of genital 
double-somite, closely set together but separate and each 
closed off by small operculum derived from sixth leg; no 
armature observed but posterior margin of operculum with 
minute spinous processes and a circular scar at the outer 
distal corner (probably indicating insertion site of long seta as 
in 5. flandricus, cfr. Fig. 5G). Copulatory pore located far 
anteriorly between genital apertures (arrowed in Fig. 4B). 
Seminal receptacles not confirmed. Paired widely separated 
secretory pores at about 2/5 distance from anterior margin. 



Anal somite (Figs. 1A-B; 4A; 5A-B) with dorsal opercu- 
lum drawn out into median, posteriorly directed, spike; 
process about as long as anal somite proper; ventral posterior 
margin spinulose medially. 

Caudal rami (Figs. 4A; 5A-B) divergent; outer distal 
corner drawn out into backwardly directed, acutely recurved, 
spinous process; with 7 setae; seta I minute, setae II and III 
located anterior to seta I, seta IV tiny and located between 
spinous process and large seta V, seta VII long and tri- 
articulate at base, seta VI minute. 

Antennule (Fig. 2A) 7-segmented, articulating on a small 
pedestal as in the male (Fig. 6B); slender, anteriorly directed 
(Fig. 1A); first segment extremely elongate, about 4 times as 
long as maximum width, with 1 short seta distally; aesthetasc 
on fourth segment fused basally to long seta; distal 2 setae of 
last segment fused basally. All setae bare; setal formula: [1, 
8, 3,2+ae, 1,2,9]. 

Antenna (Figs. 2B-D). Coxa small, not ornamented. Basis 
and proximal endopod segment fused to form allobasis, 
original segmentation marked by internal chitinous rib anteri- 
orly and incomplete suture line posteriorly near exopod; basis 
with serrate seta located on inner lateral surface; exopod 
small, 1-segmented, with 1 small, apical seta; free endopod 
articulating with allobasis at right angle (Fig. IB), lateral 
margin with 2 spines, distal margin with 1 pinnate spine and 4 
geniculate setae, the largest of which is fused basally with 
vestigial seta and bearing coarse spinule at about midway. 

Labrum (Fig. 5C) a ventrally projected, elongate, membra- 
nous outgrowth, distinctly tapering distally. Paragnaths small 
membranous lobes. 

Mandible (Figs. 2E-F) with conspicuous coxa, drawn out 
to form a slender, stylet-like gnathobase bearing small teeth 
and a long serrate seta near the apex. Palp elongate, 
2-segmented; proximal segment representing basis, slightly 
sigmoid, swollen in distal half, with 1 seta and spinular row; 
distal segment representing endopod, with 2 lateral and 3 
apical setae. 

Maxillule (Fig. 2G). Praecoxa with large, cylindrical arth- 
rite bearing 2 anterior surface setae and 6 setae along the 
distal margin; coxal endite with 2 setae; palp representing 
fused basis and rami; exopod, endopod, proximal and distal 
basal endites represented by 1, 2, 2 and 3 setae, respectively. 

Maxilla (Fig. 2H) reduced, 2-segmented. Syncoxa with 
single endite bearing unipinnate seta and conspicuous 
aesthetasc-like structure representing modified seta with chi- 
tinized dorsal margin and tubular membranous part ventrally; 
exit of maxillary gland discernible in proximal half. Allobasis 
drawn out into pinnate claw bearing serrate seta at its base. 
No trace of endopod. 

Maxilliped (Fig. 4F) subchelate. Syncoxa and basis without 
armature but with 3 spinular rows each. Endopod repre- 
sented by strong claw bearing tiny spinules along distal half of 
inner margin; an accessory setule is located at the base of the 
claw. 

PI (Fig. 3A). Praecoxa a small sclerite located around the 
outer lateral margin of the limb base. Intercoxal sclerite a 
minute rounded plate. Coxa with spinular row. Basis with 
inner and outer basal seta and with spinules at middle distal 
margin. Exopod 2-segmented, proximal segment with blunt 
spine bearing long setules, distal segment with 2 geniculate 
setae and 1 unipinnate spine. Endopod 2-segmented, elon- 
gate, prehensile; proximal segment about twice as long as 
exopod, with serrate inner seta near proximal margin; distal 
segment with 1 geniculate seta, 1 short claw and a patch of 



86 



R. HUYS AND S. OHTSUKA 




Fig. 1 Syrticola intermedius sp. nov. female. A, Habitus, dorsal; B, same, lateral. 



NEW SPECIES OF SYRTICOLA FROM JAPAN 

20 jj 



87 




g. 2 Syrticola intermedius sp. nov. female. A, Antennule; B, antenna, inner lateral view; C, distal end of antennary endopod, outer lateral 
view; D, antennary exopod, outer lateral view; G, maxillule; H, maxilla. Male. E, mandible; F, Mandible, gnathobase. 



R. HUYS AND S. OHTSUKA 






\ \\ 


\ 


i 

11 




I 

1 



Fig. 3 Syrticola intermedius sp. nov. female. A, PI, anterior; B, P2, anterior; C, P3, anterior; D, P4, anterior. 



NEW SPECIES OF SYRTICOLA FROM JAPAN 



89 




■l»iif^ 



fg. 4 Syrticola intermedins sp. nov. female. A, Urosome, ventral; B, genital double-somite, ventral; C, P5, anterior. Male. D, P5, anterior; 
E, sixth pair of legs; F, maxilliped. [Arrows in C-E indicating vestigial seta; copulatory pore arrowed in B.] 



90 



R. HUYS AND S. OHTSUKA 








■ ^:;^:-X;;:^il;m 



Fig 5 Svrftco/a mtewze^ sp. nov. male. A, Anal somite and left caudal ramus, dorsal; B, anal somite and right caudal ramus ventral- C 
labrum, anterior. Female of undescribed tantulocaridan. D, Cephalic shield, dorsal; E, same, lateral. Syrticola flandricus female F P5 
anterior; G, genital double-somite, ventral; H, anal operculum and left caudal ramus, dorsal. 




91 



ig. 6 Syrticola intermedius sp. nov. male. A, Habitus, dorsal; B, rostrum and antennule, dorsal; C, antennule, segments 3 and 4, anterior 
[armature of these segments omitted; segment 4 stippled]; D, antennulary segment 3, anterior; E, antennulary segment 4, anterior; F, 
proximal part of antennulary segment 5, anterior; G, P3 endopod, anterior. 



92 



R. HUYS AND S. OHTSUKA 



fine spinules. A distinct, ventrally directed, spinous process is 
located at the ventral midline between the maxillipedal 
syncoxae and the coxae of the first leg (Fig. 3A). 

P2-P4 (Figs. 3B-D) with 3-segmented exopods and 
1-segmented endopods. Intercoxal sclerites small, rectangu- 
lar, bare (Fig. 3C). Spines of distal exopodal segment elon- 
gate and slender in P3 and P4. Inner seta of P2 endopod 
serrate and typically recurved (Fig. 3B). Inner margin of 
endopod P3 with serrate seta and vestigial seta represented 
by setule (see inset Fig. 3C). Distal spines of endopod pinnate 
in P2-P3, bare in P4. Armature formula as follows: 





coxa 


basis 


exopod segment 


endopod segment 








1 2 


3 


1 2 


PI 


0-0 


1-1 


1-0; 1,2,0 




0-1; 0,1,1 


P2 


0-0 


1-0 


1-0; 1-0; 


1,11,0 


0,11,1 


P3 


0-0 


1-0 


1-0; 1-0; 


1,1,1 


0,1,2 


P4 


0-0 


1-0 


1-0; 1-0; 


1,1,1 


0,1,1 



Fifth legs (Figs. 4A, C) closely set together, no intercoxal 
sclerite. Baseoendopod and exopod fused to form a single 
plate with 2 secretory pores and 4 armature elements in total; 
endopodal lobe represented by long, triangular, spinous 
process without setae but with tiny spinules along proximal 
outer margin and on posterior surface; exopod presumably 
represented by weakly developed process bearing outer pin- 
nate spine, inner slender seta and a vestigial seta in between. 
Outer basal seta elongate and bare. 

Male. Body length measured from tip of rostrum to poste- 
rior margin of caudal rami (Fig. 6A) 460 \im. Ornamentation 
of body somites generally as in female; genital and first 
abdominal somites separate, with spinulose hyaline frill each. 
Sexual dimorphism in antennule, P3 endopod, P5, P6 and in 
genital segmentation. Spermatophore not observed. 

Antennule (Figs. 6B-F) indistinctly 8-segmented, articulat- 
ing on a small pedestal. Relative lengths of first two segments 
as in female. Third and fourth segment (= ancestral segment 
XIII) interdigitating as shown in Fig. 6C. Major geniculation 
between segments 6 and 7. Segmental fusion pattern: I, 
II-VIII, IX-XII, XIII, XIV-XVIII, XIX-XX, XXI-XXII, 
XXIII-XVIII. Segment 6 with 1 modified flat spine and 1 
setule, segment 7 with similar spine and 1 stubby pinnate 
element. Armature formula: [1, 9, 5, 2, 4+ae, 2, 2, 9]. 

P3 endopod (Fig. 6G) 2-segmented. Proximal segment 
unarmed. Distal segment drawn out into pinnate process 
(derived from distal spine in $) with spatulate tip bearing 2 
rows of denticles; inner margin with short pinnate seta and 
minute setule. 

P5 (Fig. 4D). Relative position, shape and armature largely 
similar to female except for the inner exopodal and outer 
basal seta being distinctly shorter. Ornamentation of endopo- 
dal lobe also slightly different with fewer spinules along the 
proximal outer margin and tiny spinules along the inner 
margin. 

Sixth pair of legs (Fig. 4E) positioned midventrally, sym- 
metrical; inner distal corner with numerous minute spinules 
and produced into a small process; armature consisting of 
inner strong spine, outer slender seta and a vestigial setule in 
between. 

Variability. An aberrant left P3 was noticed in the holo- 
type $ (Fig. 3C). 



Etymology. The species name is derived from the Latin 
inter, meaning between, and medius, meaning middle, and 
refers to the intermediate position between S. galapagoensis 
and the European species of the genus. 

Syrticola flandricus Willems & Claeys, 1982 (Figs. 
5F-H) 

Material examined. 3 5 $ from off Walcheren, The Neth- 
erlands, southern North Sea, 51° 57'25" N, 02° 40'45" E, 
depth 44.5 m, coarse sandy sediment, 08 May 1991, coll. R. 
Huys. One $ in alcohol deposited under reg. no. 1992.1077. 

The description given by Willems & Claeys (1982) is 
detailed and therefore only a few corrections to the original 
figures are noted here. 

Antenna. The exopod possesses only one seta as in S. 
intermedius and 5. galapagoensis. The oblique suture line has 
probably been mistaken for the lateral seta (compare Fig. 2D 
with Fig. 2B in Willems & Claeys (1982)), and it is conceiv- 
able that the same misinterpretation applies for S. mediterra- 
neus (cf. Willems et ai, 1987: Fig. 3A). 

Mandible. The basis bears only one seta; the supernumer- 
ary proximal 'setae' figured by Willems & Claeys are part of a 
transverse row of long spinules running around the lateral 
margin of the basis. 

Maxillule. The arthrite of the praecoxa has 6 marginal and 
2 surface setae, the coxal endite 2 setae and the distribution 
pattern of the palp setae is identical to 5. intermedius (Fig. 
2G). 

Maxilliped. The endopodal claw bears an accessory setule 
at its base. 

PI. A seta is located at the inner distal corner of the basis. 

P5. The armature of the exopodal lobe consists of an outer 
spine, an inner seta and a setule in between (Fig. 5F). 

The genital field is basically the same as in S. intermedius 
(Fig. 5G). 



REMARKS 

A single probably parthenogenetic female of a tantulocaridan 
was found attached to the pleurotergite of the P3-bearing 
somite of the holotype 2 of S. intermedius (Fig. 1A). The 
specimen is about 160 \im long and is at an early stage of 
development. The larval postcephalic trunk had been 
sloughed already but no differentiating tissue could be 
observed inside the sac. The male paratype was also infested 
by a parthenogenetic female (Fig. 6A) which was larger 
(235 jim) and attached to the pleurotergite of the genital 
somite. Inside the sac a large number of small eggs of about 
20-25 um in diameter is contained. Both tantulocaridan 
stages most likely belong to an as yet undescribed species 
which was found to infest harpacticoids belonging to at least 
two other families (Huys et al., in preparation). Since only< 
the head shield (Figs. 5D-E) is left for comparison this; 
identification has to be considered provisional. 



NEW SPECIES OF SYRTICOLA FROM JAPAN 



93 



DISCUSSION 



Syrticola intermedius is the second species to be reported 
from the Indo-Pacific, the other species (under the name (?) 
Notopontia galapagoensis) being originally described from a 
sandy beach in the Galapagos (Mielke, 1982). Both species 
resemble each other morphologically. A comparison of the 
major diagnostic characters (Table 2) reveals two species 
groups in the genus Syrticola. The European group includes 
S. trispinosus, S. flandricus and S. mediterraneus and is 
characterized by a 6-segmented antennule (segments 6 and 7 
fused) and the anal operculum possessing small spinous 
processes (Fig. 5H). The number of these projections ranges 
from (rarely) to 5, though specimens with a single small 
process have not been recorded yet (Willems et al., 1987). 
The second species group encompasses the two Indo-Pacific 
species which share a 7-segmented antennule and an opercu- 
lum drawn out into a single median strong spike. Both species 
also share the plesiomorphic 2-segmented condition of the 
male P3 endopod, but the significance of this character is 
limited since not all the males are known in the European 
species group. The zoogeographical and morphological sepa- 
ration does not warrant the upgrading of these groupings to 
generic rank, however, since S. intermedius exhibits certain 
characters found in the European species. Outgroup compar- 
ison with Notopontia and Leptopontia suggests that the 
spiniform nature of the outer exopodal spine and the loss of 
the inner baseoendopodal seta are apomorphic character 
states for the fifth legs, linking the Japanese species with its 
European congeners. The outline of the anal operculum links 
S. intermedius to (?) N. galapagoensis, justifying the latter's 
re-allocation to Syrticola by Willems et al. (1987). 

The possession of an aesthetasc-like structure on the syn- 
coxa of the maxilla in 5. intermedius is unusual. 
Re-examination of S. flandricus showed an unmodified seta 
in this position, in addition to the pinnate one also present in 
5. intermedius. Two setae are also reported on the syncoxal 
endite of 5. mediterraneus and in the outgroup taxa Notopon- 
tia and Leptopontia. The report of 3 setae on this endite in 5. 
galapagoensis (Mielke, 1982) therefore probably results from 
an misinterpretation of an aesthetasc-like structure. Without 
differential interference contrast microscopy the flaccid distal 
part is easily overlooked, thereby accentuating the lateral 
ichitinized margins as setoid structures. The relative lengths of 
the enditic 'setae' in Mielke's (1982: Abb. 18E) illustration 
are suggestive of this interpretation. 

The status of 5. trispinosus remains enigmatic as ever. A. 

Table 2 Comparison of Syrticola species. 



Scott's species is clearly closely related to 5. flandricus. 
Willems & Claeys (1982) list a number of differences but 
except for the structure of the fifth leg, all of these can be 
attributed to deficiencies in the original decription. This, 
however, does not rule out S. trispinosus as a distinct species 
since the discovery of an as yet undescribed species of 
Syrticola in the North Sea has proven species discrimination 
in this genus to be rather unreliable. Pending re-examination 
of topotypes from the Isle of Man, S. trispinosus is relegated 
to species inquirenda. 



KEY TO SPECIES 

1. Antennule 9 7-segmented, anal operculum with one large, 

median spike 2. 

Antennule 9 6-segmented, anal operculum with several small, 
spinous processes 3 . 

2. P3 endopod without inner seta; outer exopodal element P5 

setiform galapagoensis (Mielke. 1982). 

P3 endopod with inner seta; outer exopodal element P5 spini- 
form intermedius sp. nov. 

3. P5 exopodal lobe with 2 spiniform elements 

trispinosus (A. Scott, 1896). 

Only one element of P5 exopodal lobe spiniform 4. 

4. P3 endopod without inner seta 

mediterraneus Willems et al. , 1987. 

P3 endopod with inner seta 

flandricus Willems & Claeys, 1982. 



Acknowledgements. Drs Geoffrey Boxshall (The Natural History 
Museum) and Philippe Bodin (Universite de Bretagne Occidentale) 
are gratefully acknowledged for commenting on earlier drafts of the 
manuscript. We also would like to acknowledge the captain and the 
crew of the TR/V Toyoshio-maru of the Hiroshima University and 
Mr. M. Okada and Mr. Y. Endo for assistance at sea. Part of this 
study was supported by research grants of the Research Institute of 
Marine Invertebrates (1989) and the Narishige Zoological Science 
Award (1992) awarded to one of us (S.O.). R.H. is a visiting research 
fellow of the Institute of Zoology, University of Gent, Belgium. This 
is communication no. 623 of the Centre for Estuarine and Coastal 
Ecology, Yerseke, The Netherlands. 



trispinosus 



flandricus 



mediterraneus 



galapagoensis 



intermedius 



Antennule 9 

Inner seta P3 endopod 9 

Endopod P3 cf 

Duter element exopodal lobe P59o" 

Inner seta endopodal lobe P5 9 

Anal operculum processes 

;3ody length 9 (um) 
Body length cf (um) 
Distribution 



6-segmented 

? 

? 

spiniform 

absent 


6-segmented 

present 

? 

spiniform 
absent 


6-segmented 
absent 

1-segmented 

spiniform 

absent 


7-segmented 
absent 

2-segmented 
setiform 
present 


7-segmented 

present 

2-segmented 

spiniform 

absent 


several, 


several, 


several, 


one, 


one, 


small 
500 


small 
460-530 


small 
540-660 


large 
280-340 


large 
485 


? 


? 


460-500 


310-350 


460 


Irish Sea 


North Sea 


Mediterranean 


Galapagos 


Japan 



94 



R. HUYS AND S. OHTSUKA 



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Lang, K. 1948. Monographie der Harpacticiden. Hakan Ohlsson, Lund. I: 
1-896, figs. 1-361; II: 897-1682, figs. 362-607, maps 1-378. 

Mielke, W. 1982. Interstitielle Fauna von Galapagos. XXIX. Darcythompsoni- 
idae, Cylindropsyllidae (Harpacticoida). Mikrofauna Meeresbodens 87: 
1-52. 

Scott, A. 1896. Description of new and rare Copepoda. In: Report on the 
Investigations carried on in 1895 in connection with the Lancashire Sea- 
Fisheries Laboratory at University College, Liverpool. Proceedings and 
Transactions of the Liverpool Biological Society 10: 134-158 

Willems, K.A. & Claeys, D. 1982. Syrticola flandricus n. g., n. sp., a 
harpacticoid copepod from the Southern Bight of the North Sea. Crustaceana 
43: 1-8. 

, & Fiers, F. 1987. Syrticola mediterraneus n. sp., a harpacticoid 

copepod from the Bay of Calvi, Corsica. Hydrobiologia 153: 71-78. 



Bull. Nat. Hist. Mas. (Zool.) 59(1): 95 



Issued 24 June 1993 



Erratum 



The following replaces Fig. 5, Bull. Nat. Hist. Mus. (Zool.) 58(1), p. 42. 








Fig. 5. Pharyngochromis acuticeps (Steind.) Adult male; S.L. 98.0 mm. Okavango system (RUSI 34974). Photographed by Paul Skelton and 
R. Stobbs. 



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1 A new snake from St Lucia, West Indies 

6. Underwood 
11 Anatomy of the Melanonidae (Teleostei: Gadiformes), with comments on its phylogenetic 

relationships 

G.J. Howes 
33 A review of the serranochromine cichlid fish genera Pharyngochromis, Sargochromis, Serra- 

nochromis and Chetia (Teleostei: Labroidei) 

P.H. Greenwood 
45 A revision of Danielssenia Boeck and Psammis Sars with the establishment of two new 

genera Archisenia and Bathypsammis (Harpacticoida: Paranannopidae) 

R. Huys and J.M. Gee 
83 A new species of Syrticola Willems & Claeys, 1982 (Copepoda: Harpacticoida) from Japan 

with notes on the type species 

/?. Huys and S. Ohtsuka 
95 Erratum 



ZOOLOGY SERIES 

Vol. 59, No. 1, June 1993 



ISSN 0968-0470 



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Zoology Series 
ISSN 0968 - 0470 Vol. 59, No. 2, pp. 97-170 

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Printed in Great Britain at The Alden Press, Oxford 



Bull. nat. Hist. Mus. Loud. (Zool.) 59(2): 97-105 



Issued 25 November 1993 



The status of the Persian Gulf sea snake 
Hydrophis lapemoides (Gray, 1849) 
(Serpentes, Hydrophiidae) 



THE NATURAL 
HISTORY MUSEUN 

-6 DEC 1993 
ZOOLOGY L1BR AF 



ARNE REDSTED RASMUSSEN 

Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen 0, 
Denmark 



CONTENTS 



Introduction 97 

Materials and methods 97 

Systematic account 98 

Acknowledgments 103 

References 104 



Synopsis. A redescription is given of the two syntypes of Hydrophis lapemoides together with a description of 
specimens from the whole range of the species. Information on breeding and feeding biology and epizooic organisms 
of H. lapemoides is provided. Geographical variation was found between the following three areas: Andaman Sea 
and Malacca Strait, India and Sri Lanka, and Persian Gulf and Gulf of Oman. Finally the validity of H. lapemoides 
is tested against its congeners. 



INTRODUCTION 



MATERIALS AND METHODS 



In 1849, Gray described Hydrophis lapemoides on the basis of 
two specimens from Sri Lanka and Madras, respectively. In 
addition to the syntypes Smith (1926) had eight specimens 
available for characterizing the species. Smith recorded H. 
lapemoides from the Persian Gulf, and coasts of India and 
Ceylon, and considered it as a rare species. Vols0e (1939) 
collected eight specimens in the Persian Gulf and the Gulf of 
Oman, and concluded that H. lapemoides is a fairly common 
snake in these waters. Minton (1966) mentioned two speci- 
mens from the coast of Pakistan, and in 1981 Toriba & Sawai 
extended the known range of H. lapemoides from the East 
coast of India to Penang Island, Malaysia. Tamiya et al. 
(1983) identified some sea snakes from the Philippines as H. 
lapemoides, the identification, however, was questioned by 
Rasmussen (1987). Rasmussen recorded specimens from Sin- 
gapore and Phuket Island, Thailand, thereby confirming the 
presence of H. lapemoides in the Malacca Strait and Anda- 
man Sea, respectively. Recently Gasperetti (1988) considered 
H. lapemoides as the most frequent sea snake of the Persian 
Gulf. 

On the basis of my own collections in 1985, 1987 and 1989 
from Phuket Harbour, and in 1990 from Bahrain, Persian 
Gulf, I have a most welcome opportunity to describe H. 
lapemoides from its whole range, and to test the validity of H. 
lapemoides against its congeners. Some comments on the 
biology of H. lapemoides are also given. 



Material examined 

Hydrophis lapemoides BMNH: 1946.1.7.2 (syntype) (for- 
merly III. 3. 3. a) Ceylon. 1946.1.6.91 (syntype) (formerly 
III.3.3.b) Madras. 1946.1.3.88 (type of H. stewartii) (for- 
merly 83.7.30.10) Orissa, Poorie. 1946.1.9.25 (type of H. 
holdsworthii) (formerly 72.1.26.41) Ceylon. 72.1.26.43 Cey- 
lon. 80. 1 1 . 10. 199 Gwadur, Baluchistan. 1904.6. 13.19 Mekran 
coast, Charbar. 1969.2902 Persian Gulf. 1972.689 Dubai, 
Trucial Oman. 1971.135-136 Bahrain. 1970.753 east coast of 
Bahrain. 1971.1461 Bahrain harbour. 1973.410 Sharjah, Tru- 
cial coast. 1983.1169 Najwa, Darninam reef. 1983.1163-1164 
Dammam channel. 1983.1170 Oqair Bay. 1983.1172 Half 
Moon Bay, Saudi Arabia. 1985.646 Azaiba, Batinah. FMNH: 
28310-1 1 Bahrain. 64432 Tarut Bay, Ras Tonura. 73996-97 Al 
Khobar, Arabia. 82577 Persian Gulf 26° 39'N, 50° 07'E. 
121473 Ceylon. USNM: 127993 Ras Tanura, Saudi Arabia. 
132402 Saudi Arabia. RMNH: 18026 Bahrain. ZMUC: R 
66166-173 Persian Gulf (map in Vols0e, 1939). 66101 Mal- 
acca Str. (Singapore) 1° 35'N, 103° 01'E. R 66460, 66587-603, 
66605-616, 66618-627, 66629-639 all collected from trawling 
boats in Phuket port, Phuket Island, west coast of peninsular 
Thailand. R 66927, 937, 939-941, 945, 950, 951, 954, 958, 961, 
964, 965, 967, 968, 970, 971, 973-975, 980-1001, 1004-1006, 
1009, 1011, 1013 Persian Gulf 100 km north-northeast of 
Bahrain. 



©The Natural History Museum. 1993 



98 



A. REDSTED RASMUSSEN 



mi 









J^// 
vV^ 



■ I v 



J \ \ \ H J 

• i ' /M 







'j ] + 



•yv 



'Km 




Fig. 1 Sulcate and asulcate side of everted hemipenis of H. 
lapemoides (ZMUC R 66620) from Phuket Harbour. Drawing by 
M. Andersen. 




5 mm 

Fig. 2 Lateral aspect of anterior braincase of H. lapemoides in 
which the sphenoid is included in the margin of the anterior 
orifice of cavum epiptericum. Drawing by M. Andersen. 

Methods 

The measurements and counts follow Smith (1926) with some 
alterations as described below. For lateral head scales, both 
sides of the head have been examined and numbers are given 
separately. Number of maxillary teeth is given for the right 
side only. 

Scale rows are counted directly around body (Thomas 
1976). Minimum and maximum number of rows are given for 
comparison with the counts of Smith (1926, p. XVI). 

Position of the tip of the heart and the anterior tip of the 
liver are determined in relation to the number of the adjacent 
ventral scales (VS). Relative position of the tip of the heart 
and the anterior part of the liver is expressed as the percent- 
age of the total number of ventral scales (% VS). Vertebral 
counts are obtained from soft radiographs. Three counts are 
obtained from each snake: number of body vertebrae (VB- 
body), number of tail vertebrae (VB-tail), and number of 
vertebrae from the head to the tip of the heart (VB-heart). 
Body and tail are separated by the presence of the first pair of 
forked ribs in the cloacal region; this pair of ribs is included in 
the number of tail vertebrae. Tip of the heart was indicated in 



the x-ray radiographs by inserting a needle perpendicular to 
the long axis of the body pointing at the vertebrae opposite 
the tip of the heart. Relative position of the tip of the heart is 
expressed as the percentage of the total number of body 
vertebrae (% VB). Terms and description of the hemipenis 
follow Dowling & Savage (1960). Hemipenis was described in 
everted condition. All measurements are given to the nearest 
centimeter. Meristic and mensural data given as x ±SD. 

In the following description of the two syntypes H. 
lapemoides III. 3. 3. a, a subadult female (syntype a) is given 
first, and III. 3. 3. b, a juvenile male (syntype b) is given when 
different from type a. The description of the examined 
specimens is given in parentheses when different from the 
types. 

The following are abbreviations (Leviton et al., 1985) used 
for the collections: BMNH: The Natural History Museum, 
London; FMNH: Field Museum of Natural History, Chicago; 
RMNH: Rijksmuseum van Natuurlijke Historic Leiden; 
USNM: National Museum of Natural History, Smithsonian, 
Washington; ZMUC: Zoological Museum, University of 
Copenhagen. 



SYSTEMATIC ACCOUNT 

Hydrophis lapemoides (Gray, 1849). 

Aturia lapemoides Gray, 1849:46. 

Hydrophis holdsworthii Giinther, 1872:33. 

Hydrophis stewartii Anderson, 1872:399. 

Distira lapemoides. Wall, 1909:227. 

Hydrophis lapemoides, Smith, 1926:86, 1943:461. Volsoe, 
1939:19. Minton, 1966:146. McDowell, 1972:229. Voris, 
1977:91. De Silva, 1980:399. Toriba & Sawai, 1981:134. 
Rasmussen 1987:57, 1989:413, 1992:92. Gasperetti, 1988:312. 
Bussarawit et al. 1989:222. McCarthy & Warrell, 1991:163. 

Diagnosis 

Eight to 13 maxillary teeth behind poison fang, 28-35 scale 
rows on neck, 40-57 scale rows on body. Number of ventral 
scales 288-395, tip of heart extending to ventral number 
106-155. Number of body vertebrae 164-188, tip of heart 
extending to vertebrae number 73-94. Head dark dorsally 
with curved white mark, disappearing with age. Body with 
black bands forming rhombic spots dorsally and disappearing 
with age ventrally. Tail with black bands, disappearing with 
age, posterior part normally black. 

Description of the syntypes and the examined 
specimens 

External morphological characters. Maxillary teeth 
behind poison-fang 10. Dentary teeth, pterygoid teeth and 
palatine teeth not counted on syntypes; for the examined 
specimens see Table 1. One pre- and two postoculars on both 
sides (one pre- and two or three postoculars). Three anterior 
temporals on both sides (two or three). Eight supralabials on 
both sides (7-10 in males, 7-10 in females). First and second 
supralabials in contact with nasal, second and third in contact 
with preocular, third and fourth in contact with eye, syntype 
b; only third in contact with eye, fourth is divided horizon- 
tally. Eight infralabials on both sides, first, second and third 
on each side in contact with anterior pair of sublinguals, 



THE STATUS OF HYDROPH1S LAPEMOIDES 



99 








10 



11 



12 



n 



15 



Fig. 3 Habitus of the juvenile type specimen of H. lapemoides (BMNH 1946.1.6.91) from Madras, India. Photo by G. Brovad. 



which are well developed and in contact with one another; 
third and fourth infralabials touching posterior pair of sublin- 
guals, which are well developed and separated from one 
another posteriorly. A series of small cuneated scales at the 
oral margin after the third infralabial, syntype b; second 
infralabial. Scale rows on neck 29, syntype b; 32 (28-34 in 
males, 28-35 in females), on body 45, syntype b; 51 (40-51 in 
males, 41-57 in females). Ventrals 349, syntype b; 318 (288- 
365 in males, 293-395 in females), distinct throughout, bicari- 
nate, about twice as broad as adjacent scales anteriorly, 
narrower posteriorly. Subcaudals 44, syntype b; 49 (37-56 in 
males, 36-53 in females). 

Internal morphological characters. Tip of heart 
extending to ventral scale number 127, syntype b; 119 (106- 
141 in males, 106-155 in females), %VS heart 36.38%, 
syntype b; 37.42% (34.2-41.5 in males, 33.8-40.9 in females). 
Anterior end of liver situated at ventral scale number 133, 
syntype b; 120 (110-144 in males, 107-157 in females), %VS 
liver 38.10%, syntype b; 37.74% (34.4-41.5% in males, 
34.4-41.2%). In type a, a small interval separates the heart 
and the liver. Number of body vertebrae 171, syntype b; 165 
(164-188 in males, 171-186 in females). Number of tail 



vertebrae 33, syntype b; 30 (31-40 in males, 28-38 in females). 
Tip of heart extending to vertebrae number 81, syntype b; 82 
(73-90 in males, 79-94 in females), %VB heart 47.36%, in 
syntype b; 49.70% (43.5-51.1% in males, 45.1-51.7% in 
females). 

Hemipenis. Hemipenis feebly bilobed with a bifurcate sulcus 
spermaticus (Fig. 1). Bifurcation near apical end of organ 
(Fig. 1). Organ covered with spines gradually decreasing in 
size and becoming more scattered at the distal end. A 
finger-like fold at the proximal portion opposite the sulcus 
spermaticus. 

Skull morphology (based on skulls from the Persian Gulf 
and Andaman sea (Phuket)). Posterior half of parietal with a 
distinct ridge being about 1/3 of the total length (midline). 
Supratemporals (squamosals) reach parietal, and extend as 
far posteriorly as posterior part of exoccipitals. Postorbital 
bones barely touch frontals. Ventral extensions of frontals do 
not overhang trabecular grooves. Sphenoid enters broadly 
into margin of anterior orifice of cavum epiptericum (Fig. 2). 
Sphenoid with low but distinct keel. Both anterior and 
posterior Vidian foramen on the ventral side of sphenoid, and 



100 



A. REDSTED RASMUSSEN 



Table 1 Geographic variation in number of the teeth on maxilla, 
palatine, pterygoid and dentary bone. 



n Maxillary n Palatine Pterygoid Dentary 
teeth teeth teeth teeth 



Andaman Sea 
Malacca Str. 53 

India 

Sri Lanka 5 

Persian Gulf 
Gulf of Oman 73 



10-13 20 



9-10 



8-11 



12 



8-10 



7-10 



23-30 



21-26 



20-23 



20-21 



going back, yellowish ventrally, some older specimens with- 
out a curved mark. Body with black bands (29-52) forming 
rhombic spots dorsally and disappearing with age ventrally 
(Figs. 4 and 5). Tail with black bands (5-8), disappearing with 
age, posterior part black (Fig. 4). 

Breeding biology. Six of 17 females (collected in 
September-November 1987) from Phuket were pregnant. 
Three specimens contained 2 full-term embryos each, two 
specimens contained 4 full-term embryos each, and one 
specimen contained 1 full-term embryo. Pregnant females 
were collected in the period 3rd October to 4th November. 
None of the females collected in February-March 1989 were 
pregnant. The smallest embryo measured 9 cm (3rd October) 




Fig. 4 Habitus of a juvenile and an adult H. lapemoides (ZMUC R 66992, 66993) from the Persian Gulf. Photo by M. Andersen. 



in respect of the length of the Vidian canals they are 
symmetric. Palatine exceeding maxilla in forward extension, 
and without a flange for the anterior medial process of 
maxilla. Palatine-pterygoid articulation anterior to maxilla- 
ectopterygoid articulation. Fangs separated from solid maxil- 
lary teeth by a diastema. Maxillary bone slightly longer than 
ectopterygoid. Solid maxillary teeth shorter than fangs. For 
number of teeth on maxilla, palatine, pterygoid, and dentary 
bones see Table 1. 

COLOUR. Juveniles: Head black with a yellowish curved 
mark above, body yellowish or whitish, encircled by black 
bands broadest dorsally (Figs. 3 and 4). Adults: Head dark 
dorsally with curved white mark above, starting forehead 



and the largest 26 cm (19th October). The female collected 
4th November had embryos measuring 22 cm. Thus H. 
lapemoides seems to be a k-strategist (Lemen & Voris, 1981) 
producing small clutches of relatively large offspring. 

None of the females examined from the Persian Gulf were 
pregnant, however, Vols0e (1939) mentioned three females 
with eggs, and again the clutch size was very small (two 
females with 2 eggs, one female with 3 eggs). Only two of the 
three specimens have a collection date, and both were from 
April (Vols0e, 1939). 

Feeding biology. Remains of the following four fish fami- 
lies were identified in stomach contents from H. lapemoides 
collected at Phuket harbour; Gobiidae, Labridae, Mullidae, 



THE STATUS OF HYDROPH1S LAPEMOIDES 



101 




llll|llll|ll 



10 



11 



V? 



IS 19 20 



Fig. 5 Habitus of the subadult type specimen of H. lapemoides (BMNH 1946. 1 .7.2) from Sri Lanka. Photo by G. Brovad. 



and Pseudochromidae. Pseudochromidae was found as prey 
in a sea snake stomach for the first time, and Labridae and 
Mullidae are new prey records for H. lapemoides (Voris & 
Voris, 1983). The stomach contents from H. lapemoides 
collected in Bahrain were too digested to be identified, 
however, Vols0e ( 1939) mentioned Gobiidae in stomachs of 5 
specimens of H. lapemoides from the Persian Gulf. Further- 
more Voris & Voris (1983) mentioned Anguilliformes and 
Ophicthidae as stomach contents from H. lapemoides. 

Epizooic organisms. Five of the 51 specimens examined 
from the Andaman Sea and Malacca Strait had between one 
and five barnacles (Platylepas ophiophilus) on the skin. Two 
of the seven specimens from India and Sri Lanka had three 
and 20 P. ophiophilus on the skin, respectively. 25 of the 71 
specimens from the Persian Gulf and the Gulf of Oman had 
between one and 181 P. ophiophilus on the skin. Most of the 
barnacles were on the posterior part of the body. P. ophio- 
philus is found only on sea snakes (Zann et al., 1975), and has 
been found on many species (Rasmussen, 1992; Zann, 1975). 

Distribution. H. lapemoides is found from the Persian Gulf 
in west, to the Malacca Strait (Singapore) in east. Specimens 
have been collected from the Persian Gulf, the Gulf of 
Oman, the coast of Pakistan, India, and Sri Lanka, the west 
coast of peninsular Thailand, Penang (Malaysia), and Sin- 
gapore. (Ahmed, 1975; Bussarawit et al., 1989; Gasperetti, 



1988; McCarthy & Warrell, 1991; Minton, 1966; Rasmussen, 
1987; Smith, 1926, 1943; Toriba & Sawai, 1981; Vols0e, 
1939;). 

Recent collection data. H. lapemoides was collected in 
different periods of 1985, 1987, and 1989 from fishing boats in 
Phuket harbour, Phuket Island, on the west coast of peninsu- 
lar Thailand. The most common sea snake brought to the 
harbour by fishing boats was Lapemis hardwickii (over 80% 
of all sea snakes caught by trawl) followed by H. ornatus, and 
then H. lapemoides. According to the fishermen, the sea 
snakes were caught mainly by sea-going trawler-boats, fishing 
in waters more than 10 m deep. No further information was 
available, as the fishermen were rather secretive about the 
exact position of their fishing grounds. During collection in 
the Persian Gulf (Bahrain) in February 1990, we went to an 
area about 100 km north-northeast of Bahrain, on board a 
trawling boat. On a 3 days trip we collected 7 specimens of H. 
lapemoides, 2 specimens of Thalassophina viperina, and 1 
specimen of H. ornatus. They were all caught by trawl in a 
depth of 27-30 m, the bottom material was gravel. We also 
collected sea snakes at Bahrain harbour from 6 trawling 
boats, all working in the same area as mentioned above. In a 
period of 10 days (each boat was out 3 to 4 times in that 
period), a total of 110 sea snakes was collected, and 96% of 
the snakes were identified as H. lapemoides. 



102 



A. REDSTED RASMUSSEN 



Species assignment. The material examined is separated 
into three geographical regions: Andaman Sea and Malacca 
Strait, India and Sri Lanka, and Persian Gulf and Gulf of 
Oman (Tables 2 and 3, Figs. 6 and 7). When comparing 
specimens from the three areas mentioned above, geographi- 
cal variation is found in general body form (specimens from 
the Persian Gulf look more robust than specimens from 
Andaman Sea and Malacca Strait), in scale rows on neck in 
relation to scale rows on body (Fig. 6), in number of 
vertebrae (Table 3), and in number of vertebrae in relation to 
VB heart (Table 3, Fig. 7). However, it is difficult to decide 
whether the variation indicates a cline or distinct geographic 
forms, as material is still missing from Bangladesh and 
Burma, and so are representative samples from Pakistan, 
India, and Sri Lanka. 

Both Boulenger (1896), Wall (1909) and Smith (1926) 
referred the type specimen described by Anderson in 1872 
under the name H. stewartii to H. lapemoides. Having 
examined the specimen, I have serious doubt about its 
assignment. 52 Scale rows on body in relation to 30 scale rows 
on neck (Fig. 6), and 182 vertebrae in relation to 94 VB heart 
(Fig. 7) indicate that the specimen belongs to a distinct taxon. 
But as representative material is lacking from India and Sri 
Lanka, I tentatively assign it to H. lapemoides. Further mate- 
rial may show whether it is a valid taxon. 

Dunson & Minton (1978) caught some sea snakes in the 
Philippines, during the Visayan Sea Expedition of R/V Alpha 
Helix, and identified them as H. ornatus. In 1983 Tamiya et 
al. reclassified the specimens as H. lapemoides, and later 
Rasmussen (1989) reexamined the specimens and identified 
them as H. lamberti. Comparison of the above mentioned 
specimens with H. lapemoides from Andaman Sea and Mal- 
acca Strait, shows that they differ in following characters: 
Scale rows on neck (H. lamberti, 37-45), VS heart tip (H. 
lamberti, 87-109), and VS liver (H. lamberti, 86-108), VB 
heart tip (H. lamberti, 65-71), and color pattern (Rasmussen, 
1989). Comparing the skull, H. lamberti shows a more robust 
parietal, with a longer ridge (from 1/2 to 2/3 of the total 
length of parietal bone in midline), and with a less globular 
form than H. lapemoides. A single specimen of H. lamberti 
(FMNH 313058) was collected sympatrically with H. 
lapemoides (ZMUC 66101) in the area of Singapore, and also 
here the two species are distinct on the characters mentioned 
above. 



McCarthy & Warrell (1991) referred to a specimen 
(BMNH 1987.172) from the Gulf of Siam (Samut Sakhon) as 
'//. sp near H. lapemoides'. I have examined this specimen 
and agree that it is very similar to H. lapemoides, however, it 
differs in number of scale rows around body in relation to 
scale rows on neck (Fig. 6) and in number of vertebrae in 
relation to VB heart (Fig. 7). Compared to H. lapemoides 
from Malacca Strait and Andaman Sea it is very long (1.1m) 
and very robust in body and head form. In general shape it is 
much closer to H. ornatus and H. lamberti, although the 
characters differ here, too. Accordingly I think H. sp. near 
H. lapemoides' should be separated from H. lapemoides, but 
further studies are needed to find out whether the specimen 
belongs to some of the more robust species in the Gulf of 
Siam or is an unknown species. 

Generic assignment 

H. lapemoides has a combination of characters which places it 
in the genus Hydrophis as defined by Smith (1926): maxillary 
bone not extending forward beyond the palatine; poison-fang 
followed, after a diastema, by from 1 to 18 teeth; palatine 
straight; nostrils superior; nasal shields in contact with one 
another; head shields large, regular; and ventrals small, 
distinct throughout and normally entire. 

McDowells subgeneric assignment. In 1972 McDowell 
recognized three subgenera in the genus Hydrophis, how- 
ever, making a cladistic analysis (Rasmussen, in press) of the 
subgenus Chitulia (formerly Aturia, see Williams & Wallach, 
1989), the results indicated that the group was paraphyletic, 
held together by plesiomorphic character states. Neverthe- 
less, many of McDowell's character states are most useful in 
making a congeneric comparison. 

Comparison with sympatric species 

In the genus Hydrophis the following species are sympatric with 
H. lapemoides: H. bituberculatiis , H. brookii, H. caerulescens , 
H. cantoris, H. cyanocinctus , H. fasciatus, H. gracilis, H. 
inornatus, H. klossi, H. lamberti, H. mamillaris, H. melano- 
soma, H. obscurus, H. ornatus, H. spiralis, H. stricticollis, and 
H. torquatus. (Bussarawit et al., 1989; De Silva, 1980; Gasper- 
etti, 1988; McCarthy & Warrell, 1991; Minton, 1966; Murthy, 



Table 2 Geographic variation of external and internal characters in H. lapemoides. 





Sex 


n 


Ventrals 


VS-heart 


% VS-heart 


VS-liver 


% VS-liver 


Andaman 


M 


28 


288-347 


106-131 


34.2-40.6 


110-133 


35.7-41.3 


Sea and 




jc±SD 


317±13 


118±6.7 


37.5±1.4 


120±6.2 


38.1 + 1.3 


Malacca 


F 


23 


299-378 


106-140 


33.8-38.5 


107-143 


34.4-39.1 


Str. 




Jt+SD 


341 ±20 


122±7.7 


35.7±1.2 


124+7.7 


36.3+1.2 


India 


M 


2 


313-318 


114-119 


36.4-37.4 


114-120 


34.4-37.7 


and Sri 




x±SD 


315±3.5 


116±35 


36.9±0.7 


117+4.2 


37.1+0.9 


Lanka 


F 


5 


313-376 


117-145 


35.1-39.0 


117-146 


35.1-39.3 






x±SD 


347±24 


127±12 


37.3+1.8 


129+13 


37.8±1.8 


Persian 


M 


45 


293-369 


111-141 


35.0-41.5 


113-144 


36.3-41.5 


Gulf and 




x±SD 


320±16 


123±7.2 


38.5+1.4 


124+7.1 


38.8±1.4 


Gulf of 


F 


25 


300-395 


114-155 


34.6-40.9 


114-157 


34.6-41.2 


Oman 




x±SD 


347±23 


129±10 


37.5+1.6 


130+10 


37.7±1.6 



VS-heart, VS-liver = position of tip of the heart and anterior tip of liver in relation to the number of the adjacent ventral scales, respectively. %VS-heart, 

% VS-liver = relative position of tip of the heart and anterior tip of the liver in number of ventral scales, expressed as percentage of total number of ventral scales. 



THE STATUS OF HYDRO PHIS LAPEMOIDES 

Table 3 Geographic variation of internal characters in H. lapemoides . 



103 





Sex 


/; 


VB-body 


VB-heart 


%VB-heart 


VB-tail 


Andaman 


M 


28 


164-174 


73-83 


43.5-48.0 


31-38 


Sea and 




x±SD 


170±2.7 


79±2.3 


46.7±1.1 


34±2.0 


Malacca 


F 


23 


171-180 


79-86 


45.1-49.1 


28-38 


Str. 




*±SD 


174±2.5 


82±1.8 


47.0±1.1 


31±2.5 


India 


M 


2 


165-174 


79-82 


45.4-49.7 


37(/?=l) 


and Sri 




,r±SD 


169±6.4 


80±2.1 


47.5±3.0 




Lanka 


F 


5 


171-182 


81-94 


45.8-51.7 


30-35 






,r±SD 


176±4.9 


85±5.1 


48.5±2.3 


33±1.9 


Persian 


M 


45 


171-188 


79-90 


45.6-51.1 


33-40 


Gulf and 




±SD 


177±3.3 


85±2.3 


47.9±1.3 


37±1.7 


Gulf of 


F 


26 


172-186 


81-90 


46.6-49.2 


30-36 


Oman 




,v±SD 


181+3.6 


86±2.4 


48.0±0.8 


34±1.7 



VB-body = number of body vertebrae. VB-heart = position of the tip of (he heart in relation to the number of vertebrae. % VB-heart = relative position of tip of 
the heart in number of vertebrae, expressed as percentage of total number of vertebrae. VB-tail = number of tail vertebrae 



1985; Rasmussen, 1987, 1989, 1992; Smith, 1926, 1930, 1943; 
Taylor, 1965; Toriba & Sawai, 1981; Tweedie, 1983). 

The sympatric species differ from H. lapemoides in the 
following characters: H. cyanocinctus and H. spiralis have the 
sphenoid nearly excluded from the ventral margin of the optic 
fenestra (McDowell, 1972; Rasmussen, 1992: Fig. 5), a lesser 
number (< 9) of maxillary teeth, (< 20) pterygoid teeth, and 
(< 20) dentary teeth, and a different colour pattern (Bussa- 
rawit et al., 1989; McDowell, 1972; Rasmussen, in press; 
Smith, 1926). H. brookii, H. cantoris, H. fasciatus, H. 
gracilis, H. klossi, H. melanosoma, and H. obscurus have a 
triangular flange on the palatine (McDowell, 1972; Rasmus- 
sen, 1992: Fig. 4), and a lesser number (< 8) of maxillary 
teeth, (< 17) pterygoid teeth, and (< 16) dentary teeth 
(McDowell, 1972). H. bituberculatus has a lesser number 
(25-29) of scale rows around neck, a lesser number (247-290) 
of ventrals, a lower position (90-105 VS) of heart tip, and a 
different colour pattern (Rasmussen, 1992). H. caerulescens 
has a higher number (14-18) of maxillary teeth (Smith, 1926), 
a higher position (96-99 VB, based on 3 specimens from 
Phuket harbour) of heart tip, and a different colour pattern 
(Bussarawit et al., 1989; Smith, 1926; Tweddie, 1983). H. 
inornatus (type specimen BMNH 1946,1.1.27 formerly 
III. 7.1. a.) has a lesser number (253) of ventrals, a lower 
position (86 VS) of heart tip, a lower position (67 VB) of 
heart tip, and a different colour pattern (Rasmussen, 1989). 
H. lamberti is compared with H. lapemoides in the section 
concerning species assignment. H. mamillaris has a smaller 
head, a lesser number (25-29, 35-43) of scale rows on neck 
and body, and a different colour pattern (Minton, 1966; 
Smith, 1943). H. ornatus has a lesser number (224-294) of 
ventrals, a lower position (72-104 VS) of heart tip, a lower 
position (59-65 VB) of heart tip, and a different colour 
pattern (Rasmussen, 1989). H. stricticollis has a smaller head, 
a higher number (> 200 VB, Voris, 1975, and own observa- 
tion) of vertebrae, and the hemipenis is bilobed half way 
down. H. torquatus has a higher position (91-105 VB) of 
heart tip, and a lesser number (7-8) of maxillary teeth (only in 
Malacca strait) (own observation). 

Comparison with allopatric species 

In the genus Hydrophis the following species are allopatric 
with H. lapemoides: H. belcheri, H. coggeri, H. czeblukovi. 



H. elegans, H. geometricus, H. macdowelli, H. melanoceph- 
alus, H. pacificus, H. parviceps, and H. vorisi. (Bussarawit et 
al., 1989; Cogger, 1975; Kharin, 1983, 1984a. 1984b; McCar- 
thy & Warrell, 1991; Smith, 1986; Smith, 1926, 1930, 1935). 
The allopatric species differ from H. lapemoides in the 
following characters: H. coggeri, H. czeblukovi, H. elegans, 
H. melanocephalus, and H. pacificus have the sphenoid 
nearly excluded from the ventral margin of the optic fenestra 
(Kharin, 1984b; McDowell, 1972; Rasmussen, 1992: Fig. 5) 
and a lesser number (< 9) of maxillary teeth, (< 20) 
pterygoid teeth, and (< 20) dentary teeth (Kharin, 1984b; 
McDowell, 1972). H. parviceps and H. vorisi have a triangu- 
lar flange on the palatine (Kharin, 1984a; McDowell, 1972; 
Rasmussen, 1992: Fig. 4), and a lesser number (< 8) of 
maxillary teeth, (< 17) pterygoid teeth, and (< 17) dentary 
teeth (Kharin, 1984a; McDowell, 1972; Smith, 1935). H. 
belcheri has a lesser number (24-26, 32-36) of scale rows on 
neck and body, a lesser number (14-17) of pterygoid teeth, 
and no cuneate scales at infralabials (McCarthy & Warrell, 
1991). H. geometricus has a high number (51-58, a small 
overlap) of scale rows on body, and a different colour pattern 
(Smith, 1986:152 Fig. 1). H. macdowelli has a lesser number 
(< 8) of maxillary teeth, (< 16) pterygoid teeth, and (< 17) 
dentary teeth, and a lesser number (256-266) of ventrals 
(Kharin, 1983). 



Acknowledgements. I thank the staff of The Phuket Marine Bio- 
logical Center, Thailand, CODEC Project, Chittagong, Bangladesh, 
Ministry of Commerce and Agriculture, Directorate of Fisheries, 
Bahrain, S. Bagge, Cowi-Almoayed Gulf, Bahrain, J. Jensen, 
Danida, and M. Andersen who helped me during the collection. The 
Natural History Museum, London, Field Museum of Natural His- 
tory, Chicago, Rijksmuseum van Natuurlijke Historie, Leiden, 
National Museum of Natural History, Smithsonian, Washington for 
loan of specimens. M. Andersen, A.B Helwigh, and especially Dr. 
C. McCarthy (BMNH) and Dr. J. B. Rasmussen (ZMUC) for 
valuable advice and constructive criticism of the manuscript. The 
study was supported by Dansk Naturhistorisk Forening, The 
Johannes Schmidts Grant, The Krista and Viggo Petersens Grant, 
The Danish Research Academy, and The Danish National Research 
Council, Grant no. 11-8209. 



104 



A. REDSTED RASMUSSEN 



38 



36 



34 



32 



w 30 



28 



26 



38 



+ Andaman sea and Malacca Str. 
D India and Sri Lanka 

Persian Gulf and Gulf of Oman 

+ + 

+ + + + 

• ■ + + + + + ffl 
x x - + --* + + 

X X X X >. D X 
X X X > 
X > • 



40 



42 



44 46 48 
Scale rows on body 



50 



52 



54 



92 
90 
88 
86 
84 
82 
80 
78 
76 
74 
72 
70 



■ • 


• 


X • > 




... 


XX 


. . 




. _ 




* + XX> 




+ + + ■ 




+ + 




+ * D 





+ + 



+ Andaman Sea and Malacca Str. 
□ India and Sri Lanka 

Persian Gulf and Gulf of Oman 



162 166 170 174 178 182 
Number of vertebrae 



186 



190 



40 -, 


+ 

D 


Andaman sea and Malacca Str. 
India and Sri Lanka 


38 




Persian Gulf and Gulf of Oman 
H. sp. near H. lapemoides 


36- 




+ + + + + + 


34 




O ■ + + 

+ m + + 


32 




. . . + . + 

X + ■ ■ 


30 


X 


U*-Type of tL slewadi 

□ 


28 

?6 




X 



96 

94 
92 
90 
88 
86 
84 
82 
80 
78 



38 40 42 44 46 48 50 52 54 56 58 60 
Scale rows on body 



76 



+ 


Andaman Sea and Malacca Str. 




□ 


India and Sri Lanka 








Persian Gulf and Gulf of Oman 


'. *-Type of tL siewani 


o 


tL sp. near hL lapemoides 




X 






■ 


X X 






■ 


■ ■ 




■ ■ 




< X X X 




□ + 


IEI 






■ + 


■ • 






+ - + 


+ 






+ + + + + • 








+ + + + D 








D * + + + + 








+ + 








+ 







"168 170 172 174 176 178 180 182 184 186 188 
Number of vertebrae 



Fig. 6 Relation between number of scale rows on body and 
number of scale rows on neck in males (top) and females 
(bottom) of H. lapemoides, showing geographic variation. 



Fig. 7 Relation between number of body vertebrae and position of 
heart tip in males (top) and females (bottom) of H. lapemoides, 
showing geographic variation. 



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Bull. not. Hist. Mus. Lond. (Zool.) 59(2): 107-124 



Issued 25 November 1993 



Taxonomic revision of some Recent 
agglutinated foraminifera from the Malay 
Archipelago, in the Millett Collection, The 
Natural History Museum, London 

P. BRONNIMANN t 

9G, Chemin de Bedex, 1226 ThonexIGeneva, Switzerland 

J.E.WHITTAKER 

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5 BD 

Synopsis. Eleven species of Recent agglutinated foraminifera in the Millett Collection from the Malay Archipelago, 
have been re-examined and revised systematically. They were originally described and illustrated in 1899 and 1900 
with excellent lithographic drawings in the hand of Millett. With rare exceptions, the actual figured specimens, 
though not marked as such, have been recognised in his Collection. The species are here redescribed, re-illustrated 
by SEM photography and, where necessary, supplemented by new data, notably from similar environments in 
Brazil. All, with the exception of Paratrochammina simplissima (Cushman & McCulloch) and possibly Trocluim- 
mina? milletti sp.nov., are brackish-water forms. 

New taxa arc Trunculocavus durrandi gen. et sp.nov. and Trochammina? milletti sp.nov. A lectotype is designated 
for Acupeina triperforata (Millett), while Ammobaculites salsus var. distinctus Cushman & Bronnimann is elevated 
to specific rank and placed in Ammotium. All the species reviewed in this paper belong to the Suborder 
Trochamminina. 



INTRODUCTION 



Durrand (1898) gives what locality information there is about 
the Malay samples from which Fortescue William Millett 
made his classic study of the foraminifera. Durrand had for 
several years, out of his own interest, obtained small samples 
from around the SW Pacific and had examined them for the 
microscopical fauna and flora. In 1889 he had succeeded in 
getting the Netherlands India Steam Navigation Company 
(then controlled by the British India Steam Navigation Com- 
pany) ... 'to instruct the commanders of their fleet plying 
about the islands of the Archipelago, to collect bottom from 
each port of call'. The 'cleaned material' was picked over first 
by Durrand and then the foraminifera were determined by 
Millett and published (1898-1904) in 17 parts in the Journal 
of the Royal Microscopical Society. In all, 468 species and 
varieties were listed by Millett, 45 of them new. The descrip- 
tions were accompanied by 19 plates of quite exceptional and 
accurate drawings, from Millett's own hand. 

The samples came from anchor mud where the ships were 
moored, more or less close inshore ... 'in about 12 or 14 
fathoms' (22-25 m). Unfortunately, a number of labels on 
the flasks of sediment became illegible through getting 
soaked by leakage, so the locality information is somewhat 
sketchy. The original samples each contained about 4 lbs 
(1.8 kg) of solid matter. 

The material came from two areas. Area 1 ('from Celebes 
in the north and west, to Java in the south and New Guinea, 



t Deceased 6.1.1993. 

©The Natural History Museum, 1993 



Aru, and the Islands in the east, including such stations as 
Banda, Amboina, Flores, Sumbawa and Timor') contains 
stations 1-16; area 2 ('Singapore in the north, Banka in the 
south, Sumatra in the west, and Borneo in the east') contains 
stations 17-31. 

As part of a major revision of shallow-water agglutinating 
foraminifera of the Indo-Pacific region (see also Bronnimann 
et al. 1992), eleven species belonging to the Trochamminina 
are here redescribed, and illustrated by scanning electron 
microscopy for the first time. The fauna has, for the most 
part, strong affinities with the foraminifera of brackish, 
mangrove sediments from other parts of the tropics, notably 
Brazil. Comparison is therefore made with material described 
by us (Bronnimann & Zaninetti, 1984a; b; Zaninetti et al., 
1977) from the mangroves of Guaratiba, Acupe and Baia de 
Sepetiba, Brazil. 

For a recent review of mangrove foraminifera in general 
and their potential for palaeoenvironmental interpretation, 
the reader is referred to an important paper by Culver (1990). 



LOCALITY INFORMATION 

Of relevance to the present revision are the following stations 
from whence the specimens came; where the name of the 
station is not mentioned, the label has become illegible. The 
sample descriptions are in Durrand's own words. 

Area 1 Station 2 [no locality]. Plastic mud, brownish 
tinted, rich in floatings. 



108 

Station 3 [no locality]. Brownish mud with lumps 
of blueish clay throughout, residue about one 
quarter-pound and floatings small. 
Station 5 [no locality]. Blue ooze, residue and 
floatings small. 

Station 9 [no locality]. Results poor. 
Station 11 [no locality]. 
Station 12 [no locality]. 

Station 14. Similar to Station 13 [Segaar, New 
Guinea, coral sand and mud, residue about six 
ounces, floatings rich]. 
Station 15 [no locality]. 
Area 2 Station 17. Muntok Banka, blue mud, residue 
eight ounces, floatings rich. 

Station 19 [no locality]. Earthy coloured, river- 
looking mud, few foraminifera. 
Station 21. Paney, northeast coast of Sumatra. 
Station 27 [no locality]. 
Station 28 [no locality]. 

Durrand (1898:257) adds a postscript, stating that ... 'it is 
important to bear in mind all this series was obtained from 
shallow water close inshore . . .'. It is clear from the aggluti- 
nating foraminifera revised here, that most of the localities 
were in fact brackish, associated with mangroves. 



SYSTEMATIC DESCRIPTIONS 



Order FORAMINIFERIDA Eichwald, 1830 

Suborder TROCHAMMININA Bronnimann & Whittaker, 

1988 

Apart from the hierarchy listed above, no further suprage- 
neric taxa will be used. Until we can be certain that the 
families and superfamilies of agglutinating foraminifera used 
by Loeblich & Tappan (1987) represent homogeneous units 
with respect to the wall structure, then it is better, for the 
present, not to use them. Similarly, genera are used in 
'inverted commas' when the wall structure of their type 
species has not yet been examined. The eleven species 
described here, at least, all have a Trochamminina-type wall, 
defined by Bronnimann & Whittaker (1988) as . . . 'consist- 
ing of organic and agglutinated phases. Agglutinant bound by 
organic cement and outer and inner organic sheets. Devoid of 
perforations or alveolar pseudopores'. 

The synonymies are not meant to be comprehensive, they 
are selective, merely listing the original reference, junior 
synonyms, changes of generic combination and important 
citations from the study area. 



P. BRONNIMANN AND J.E. WHITTAKER 
Genus ACUPEINA Bronnimann & Zaninetti, 19846 

Type species. Haplophragmium salsum Cushman & Bronni- 
mann, 1948a (= junior subjective synonym of Haplophrag- 
mium agglutinans d'Orbigny var. triperforata Millett, 1899). 

Acupeina triperforata (Millett, 1899) Figs 1.2, 13-15 

1899 Haplophragmium agglutinans d'Orbigny var. triper- 
forata Millett: 358{pars); pi. 5, figs 2a, b (lectotype) 
only; non figs 3a, b. 

1948a Haplophragmium salsum Cushman & Bronnimann: 
16,17; pi. 3, figs 10-13. 

1965 Lituola salsa (Cushman & Bronnimann); Bronni- 
mann & Zaninetti: 608-615; figs 1-3. 

19846 Acupeina salsa (Cushman & Bronnimann); Bronni- 
mann & Zaninetti: 219-222; figs Al-4, Bl,2. 

19846 Acupeina triperforata (Millett); Bronnimann & 
Zaninetti: 222 (addendum). 

1988 Acupeina triperforata (Millett); Bronnimann & Whit- 
taker: 112; pi. 4, figs 1-7. 

Remarks. Millett (1899, pi. 5, figs 2,3; here reproduced as 
Figs 1.2, 3) illustrated four views of his new variety triperfo- 
rata. Examination of the original material shows that two 
different brackish species are involved: Acupeina triperforata 
(Millett) and Arenoparrella mexicana (Kornfeld). 

The individual drawn by Millett (1899, pi. 5, figs 2a, b; here 
reproduced in Fig. 1.2a,b) in side and apertural views, has 
been re-illustrated by SEM in Figs 13-15. The micrographs 
show side and edge views of a test, initially streptospiral then 
uniserial, and the detail of the multiple aperture which 
consists of three closely spaced, virtually equidistant pores (of 
around 25 um diameter) with upturned rims. Millett appar- 
ently believed that the aperture of his new variety invariably 
consisted of the three rounded pores, hence the name. The 
individual in Figs 13-15 is undoubtedly Millett's figured 
specimen and is here formally designated lectotype. 

The specimen drawn by Millett (1899, pi. 5, figs 3a, b; 
reproduced here in Fig, 1.3a,b) in side and apertural views, 
has been re-illustrated by SEM in Figs 9-12 not only to show 
both sides of the test but the details of the composite 
aperture. Its morphology is quite different from the lectotype 
of H. agglutinans var. triperforata. It represents, in fact, a 
typical specimen of Arenoparrella mexicana (Kornfeld, 
1931)(see below). It is unfortunate that Loeblich & Tappan 
(1987: 21, pi. 71, figs 3,4) illustrated this very specimen, 
together with the lectotype, as A. triperforata. It is also worth 
noting that Millett's pi. 5, fig. 3b is the edge view of fig. 3a, 
but as can be seen from our SEM illustration, rather mislead- 
ing. It purports to show only three large pores with everted 
borders. In reality, it has a single oblique-perpendicular slit 
and 12 small peripheral pores of 5-6 um diameter, devoid of 
rims. Closer examination of Millett's apertural view (see Fig. 
1.3b) may just show the termination of the slit (the specimen 
is tilted forward), but the determination of the pores is still 
seriously in error. 



Fig. 1.1-1.10, 1.12 Reproduction of part of Plate 5 of Millett (1899). The original identifications were as follows: Fig. 1.1, Haplophragmium 
agglutinans (d'Orbigny), X112; Fig. 1.2, 3, H. agglutinans var. triperforata var.nov., X112; Fig. 1.4-6, H. cassis (Parker). X112; Fig. 1.7, 
H. cassis (Parker) or ?Reophax, X75; Fig. 1.8, H. compressum Goes, X75; Fig. 1.9, H. nanum Brady, X112; Fig. 1.10, H. anceps Brady, 
X56; Fig. 1.12, Trochammina ochracea (Williamson), X75. Reproduced by permission of the Royal Microscopical Society. 

Fig. 2.1 Reproduction of part of Plate 1 of Millett (1900). It was originally identified as Bigenerina digitata d'Orbigny var., X169. 
Reproduced by permission of the Royal Microscopical Society. 



TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 



109 




110 



P.BRONNIMANN AND J.E. WHITTAKER 



Bronnimann & Zaninetti (1984b: 222, Addendum) have 
shown that Haplophragmium agglutinans d'Orbigny var. trip- 
erforata Millett (1899) is identical with H.salsum Cushman & 
Bronnimann (1948a), which is the type species of Acupeina 
Bronnimann & Zaninetti, 19846. 

Lectotype. The individual illustrated by Millett (1899, pi. 5, 
figs 2a, b; Figs 1.2, 13-15 herein) is designated lectotype of//. 
agglutinans var. triperforata , now Acupeina triperforata. It is 
deposited in the collections of the BMNH, no. 
1955.11.1.1076. 

Dimensions (lectotype). Height of test — 380 um; diam- 
eter of coiled portion — 235 urn; maximum diameters of 
apertural pores — 25 um, with everted rims 4 urn high. 

Environment. This species ... 'is not uncommon at Station 
9, and occurs also, but very sparingly, at Station 5' of Area 1. 
At Station 9, Millett (1899: 359) also reported Haplophrag- 
mium cassis (Parker) (= Ammoastuta salsa Cushman & 
Bronnimann and Ammotium spp.), all brackish water spe- 
cies. Both Acuipeina triperforata and Arenoparella mexicana 
are also exclusively brackish forms, occurring commonly in 
tropical to subtropical mangrove swamp sediments. 



Genus AMMOSASTUTA Loeblich & Tappan, 1984 

Type species. Ammoastuta salsa Cushman & Bronnimann, 
1948a. 



Ammoastuta salsa Cushman & Bronnimann, 194&V* 

Figs 1.6,35 



1899 



1948a 

1970 
1986 



Haplophragmium cassis (Parker); Millett (pars): 359; 
pi. 5, figs 6a, b only (non figs 4,5,7) (non Lituola 
cassis Parker, 1870). 

Ammoastuta salsa Cushman & Bronnimann: 17; pi. 
3, figs 14-16. 

Ammoastuta salsa Cushman (sic); Hofker: 3. 
Ammoastuta salsa Cushman & Bronnimann; Bronni- 
mann: 29-AA; figs 1-7. (a. v. for synonymy). 



Remarks. Millett (1899: 359, pi. 5, figs 6a,b; here repro- 
duced as Fig. 1.6a,b) figured side and edge views of a slightly 
damaged, but clearly recognizable specimen of Ammoastuta 
salsa under the name of Haplophragmium cassis (Parker). He 
also illustrated two different species of Ammotium (pi. 5, figs 
4a,b, 5a, b; Figs 1.4, 5) and, used for all these different 
morphologies the same name, as he thought . . . 'the Malay 
specimens of this species [//. cassis] are very variable in form, 
some of them being extremely compressed, and composed of 
numerous chambers'. 

The SEM photograph of the side view (Fig. 35), although 
now slightly more damaged, is demonstrably of the same 
specimen as in Millett's drawing. The tight initial coil cannot 
be seen, but on the other hand, the final two chambers of the 



juvenile stage are clearly visible. The adult consists of at least 
7 elongate uniserial chambers which make up the main 
portion of the compressed test. 

Bronnimann's (1986) morphological revision of A. salsa 
has shown that the test starts with a tight early spiral 
consisting only of a proloculus and deuteroloculus. On the 
basis of this arrangement, Ammosastuta is correctly placed in 
the Lituolidae. Loeblich & Tappan (1987: 79) accepted this 
interpretation, but stated that the second chamber is growing 
in the . . . 'opposite direction' (without saying in respect to 
what). This is simply not the case. The second chamber 
develops from a porus in the side of the proloculus. It is just 
the normal forward continuation, considering the flow of the 
protoplasm, which produces the elongate deuteroloculus with 
a porus at its apex. Hence the embryonic chambers form a 
tight, reduced spiral (see Bronnimann, 1986: 32, fig. 3). 

Ammoastuta salsa is occasionally placed in synonymy with 
Ammobaculites (=Ammoastuta) ineptus Cushman & McCul- 
loch, 1939. Cushman & Bronnimann (1948a) regarded the 
two as distinct, as did Bronnimann (1986). An examination 
by Bronnimann of the two paratypes of A. ineptus, deposited 
in the collections of the U.S. National Museum of Natural 
History, Washington, confirms this separation. Of the 
paratypes, only one, registration no. 35826, is well preserved. 
It is definitely an Ammoastuta, but differs from the com- 
pressed A. salsa by having a strongly inflated test. 

Dimensions of figured specimen (BMNH no. 
1955.11.1.1121). Maximum height (damaged) — 280 um. 

Environment. Recorded from Station 9, Area 1. It occurs 
together with Acupeina triperforata, Ammotium spp. and 
Arenoparella mexicana, all typical brackish water species. 



Genus AMMOBACULITES Cushman, 1910 

Type species. Spirolina agglutinans d'Orbigny, 1846. Lecto- 
type designated by Loeblich & Tappan (1964: C241, figs 
251.6a,b). 

Remarks. The genus Ammobaculites Cushman (1910) con- 
tains free agglutinated tests with a simple interior; the early 
portion is planispiral, the later part uncoiled and rectilinear. 
It is radially-symmetrical in transverse section. The single 
aperture is terminal, areal and radially symmetrical. The wall 
structure of the type species is unknown. 

This definition is more restrictive than Loeblich & Tap- 
pan's (1987: 74) as it not only excludes streptospiral and 
trochospiral initial coils, but also laterally compressed tests. 
The transverse sections of the chambers of the uncoiled 
portion of the test and the outlines of the terminal apertures 
are radially symmetrical; these features are regarded as 
important generic criteria. 

The wall structure of Ammobaculites exiguus, the species in 
the Millett Collection, is of the Trochamminina type. If A. 



Figs 3-8 Trunculocavus durrandi gen. et sp.nov. Figs 3,4, Detail of aperture (X900) and side view (X160), respectively. Holotype, BMNH 

no. 1955.11.1.187; Fig. 5, Side view (X175). Paratype, BMNH no. 1911.11.1.189; Figs 6-8, Detail of initial coil (X540), aperture (X730) 

and side view (X160), respectively. Paratype, BMNH no. 1955.11.1.188. 
Figs 9-12 Arenoparrella mexicana (Kornfeld). Detail of apertures (X480), side, edge and view of other side (X160), respectively. BMNH 

no. 1955.11.1.1075. 
Figs 13-15 Acupeina triperforata (Millett). Edge and side view (X160) and detail of aperture (X700), respectively. BMNH no. 

1911.1. .1.1076. 
All from the Millett Collection, Malay Archipelago. 



TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 



111 




112 



P. BRONNIMANN AND J.E. WHITTAKER 



agglutinans, the type species, should have the same wall type, 
then exiguus would be correctly placed in Ammob acuities. If 
not, then it would have to be placed in a new genus. In view 
of these uncertainties, Ammobaculites is placed in inverted 
commas in our treatment of 'A '. exiguus. 

'Ammobaculites' exiguus Cushman & Bronnimann, 
19486 Figs 1.1,42-44 

1885 Haplophragmium agglutinans (d'Orbigny); Balkwill 
& Wright: 330; pi. 13, figs 18?, 19,20 (non d'Orbigny, 
1846). 

1899 Haplophragmium agglutinans (d'Orbigny); Millett: 
357, pi. 5, figs la,b. 

1938 Ammobaculites agglutinans (d'Orbigny); Barten- 
stein:391;fig. 14. 

19486 Ammobaculites exiguus Cushman & Bronnimann: 
38; pi. 7, figs 7,8. 

19526 Ammobaculites cf. exiguus Cushman & Bronnimann; 
Parker: 443 ;pl. 1, figs 16,17. 

1952 Ammobaculites agglutinans (d'Orbigny); Rottgardt: 
180; pi. 1, fig. 4. 1954 Ammobaculites exiguus Cush- 
man & Bronnimann; Phleger: 633; pi. 1, fig. 5. 

1956 Ammobaculites sp. B, Warren: 139; pi. 1, figs 22-24. 

1957 Ammobaculites exiguus Cushman & Bronnimann; 
Todd & Bronnimann: 23; pi. 2, fig. 7. 

1962 Ammobaculites exiguus Cushman & Bronnimann; 
Benda & Puri: 335; pi. 1, fig. 15. 1973 Ammobacu- 
lites balkwilli Haynes: 25-27; pi. 2, figs 2,3; pi. 29, 
figs 5,6; text-fig. 4.1-5. 

1978 Ammobaculites dilitatus (sic) Cushman & Bronni- 
mann; Schafer & Cole: pi. 3, fig. 9 (non Cushman & 
Bronnimann, 19486). 

1980 Ammobaculites dilatatus Cushman & Bronnimann; 
Scott & Medioli: 35; pi. 1, figs 9,10. 

71983 Ammobaculites exiguus Cushman & Bronnimann; 
Haman: 72; pi. 5, figs 1^1. 

1983 Ammobaculites diversus Cushman & Bronnimann; 
Haman: 72; pi. 4, figs 14,15 (non Cushman & Bron- 
nimann, 19486). 

1986 Ammobaculites exiguus Cushman & Bronnimann; 
Bronnimann & Keij: pi. 3, fig. 7. 

Remarks. Millett (1899: pi. 5, figs la,b; here reproduced as 
Figs l.la,b) illustrated, without description, a typical speci- 
men of exiguus under the name of Haplophragmium agglutin- 
ans (d'Orbigny). The same specimen (BMNH no. 
1955.11.1.1057) is re-illustrated by SEM in our Figs 43,44. 
The oblique view (Fig. 43) shows the radially-symmetrical 
areal and terminal aperture, which is larger than in Millett's 
drawing. It is not bordered by a rim as that shown by 
Haman's (1983, pi. 5, figs 1-4) 'A. exiguus', which may 
represent a different species. Millett's specimen has four 
uniserial chambers which follow from a planispiral, tightly 
enrolled early test. The agglutinant is coarse and the sutures 
in the initial portion are not well defined; on the uniserial 
portion, they are distinct, however, and run perpendicularly 
to the elongate axis of the test. 

Illustrated in Fig. 42 (BMNH no. 1911.11.1.1058) is a 
smaller, albeit damaged specimen, which is more typical of 
the size of the Malay material. Four radial sutures can be 
recognized in the coiled portion and there are three chambers 
in the uniserial part; the final chamber is crushed. 

One of us (P.B.) has re-examined the holotype of A. 



exiguus (registration no. 56761) in the U.S. National Museum 
of Natural History. Its overall morphology corresponds well 
with Millett's illustrated specimen of H. agglutinans. How- 
ever, in its uniserial portion there are five chambers and the 
agglutinant is finer than in the Malay specimen. Nevertheless, 
the two both have a circular transverse section and a large 
radially-symmetrical, terminal aperture without a rim; the 
intercameral sutures run perpendicular to the elongate axis of 
the test, there being no suggestion of Ammotium-type 
sutures. In addition to the holotype of exiguus, there are two 
slides with paratypes: in slide no. 56762 there is a single 
paratype; under no. 56763 there are, amongst typical speci- 
mens, some very small individuals which differ from the type 
by their thin, elongate tests. These latter have also been 
encountered by us in the mangrove sediments of Acupe, 
Brazil. They represent a new species of brackish 'Ammobacu- 
lites' which will be published elsewhere. It should be noted 
that 'A', exiguus and this new, minute species, are the only 
true representatives of 'Ammobaculites' occurring in brackish 
waters. 

Dimensions of figured specimen (BMNH no. 
1911.11.1.1057). Height of test — 385 \im; diameter of initial 
planispiral portion — 135 um; diameter of final chamber — 
125 ^m; diameter of aperture — 50 \im. 

Environment. In the Millett Collection, specimens are 
labelled 'Haplophragmium agglutinans' from stations 
2,9,12,14,15,19,21 and 27; Millett notes (p. 358) that . . . 'the 
specimens are all minute, and although they occur at most of 
the Stations, are not very numerous'. According to Parker et 
al. (1953), 'A', exiguus is a species which lives in brackish as 
well as in marine waters. 



Genus AMMOTIUM Loeblich & Tappan, 1953 

TYPE SPECIES. Lituola cassis Parker (in Dawson), 1870. 

REMARKS. Ammomarginulina Wiesner, 1931 (type species: 
A. ensis Wiesner, 1931) is a deep-water genus, with a 
morphology close to that of the supposedly exclusively 
brackish-water genus, Ammotium. After having compared 
the definitions of Ammomarginulina and of Ammotium in 
Loeblich & Tappan (1987), the question arises as to whether 
the two are really synonymous. The sutures of the former are, 
however, less slanting that those of Ammotium, and the test 
is strongly compressed. Of the shape of the aperture of 
Ammomarginulina ensis nothing is known except for the fact 
that it is rounded. Small morphological differences such as 
these may not be considered sufficient to retain the two 
genera. However, they seem to represent two disparate 
homogeneous environmental groups which, should this be 
sustained, must be separated taxonomically, even if the 
morphological differences were even less pronounced (see 
also Resig's (1982: 977-978, pi. 1, figs 3-5,9) description of 
Ammomarginulina hadalensis Resig from the Peru-Chile 
Trench, depth 5846 m). Clearly, the wall structure of Ammo- 
marginulina must also be investigated. 



Ammotium morenoi (Acosta, 1940) 
1899 



Figs 32-34, 54 



Haplophragmium cassis (Parker): 359 (pars) (non 
Lituola cassis Parker, 1870). 
1940 Ammobaculites morenoi Acosta: 272; pi. 49, figs 3,8 
(holotype) only (non Fig. 1). 



TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 



113 



1948a Ammobaculites salsus Cushman & Bronnimann: 16; 

pi. 3, figs 7a, b, 8,9 (holotype figs 7a, b). 
19526 Ammoscalaria fluvialis Parker: 444; pi. 1, fig. 24 

(holotype) only (non fig. 25). 

1953 Ammobaculites salsus (et vars.) Parker et al. : 5; pi. 1, 
figs 18-25 only (non fig. 17). 

1954 Ammobaculites exilis Cushman & Bronnimann; 
Phleger: pi. 1, fig. 6 (non Cushman & Bronnimann, 
19486). 

1954 Ammobaculites salsus Cushman & Bronnimann; 

Phleger: pi. 1, fig. 7 only (non fig. 8). 
1954 Ammoscalaria fluvialis Parker; Phleger: pi. 1, fig. 11. 

1957 Ammobaculites salsus Cushman & Bronnimann; 
Todd & Bronnimann; 24, pi. 3, fig. 8 

1958 Ammobaculites salsus Cushman & Bronnimann; 
Arnal: 37; pi. 98, figs 4-7. 

1968 Ammotium salsum (Cushman & Bronnimann); 

Lutze: 25; pl.l, figs 5,6. 
1978 Ammotium salsum (Cushman & Bronnimann); Poag: 

405; pi. 5, figs 1-39. 
1980 Ammotium salsum (Cushman & Bronnimann); Scott 

&Medioli: pi. 1 , figs 11-13. 
1983 Ammotium morenoi (Acosta); Haman: 72; pi. 5, figs 

6-9. 



morphological variability. She tried to distinguish six differ- 
ent morphological types on the basis of outline, chamber 
form, and dimensions of the test and chambers. A study of 
Brodniewicz's paper, however, suggests to us that it is 
virtually impossible to separate her different morphotypes. 

Dimensions of figured specimens (MALAY SPECIMEN, 
BMNHno. 1991.11.1.1122). Height of test— 170 urn; width 
(length) — 105 urn; thickness — 35 urn. 

(BRAZILIAN SPECIMEN). Height of test — 370 um; 
maximum width — 190 um; final chamber — 225 um high; 
maximum diameter of oblong aperture — 50 um. 

Environment. Found only in Station 9 (Area 1) in associa- 
tion with Ammotium pseudocassis , A. directum, Acupeina 
triperforata, Ammoastuta salsa and Arenoparella mexicana, 
all typical brackish water species. A. morenoi is normally 
abundant in tropical and subtropical mangrove sediments but 
has also been recorded, albeit rarely, in brackish sediments of 
temperate climes (Parker, 19526; Lutze, 1968). We have 
never encountered this species in the British Isles or in the 
Mediterranean. 

Observations on certain synonyms and non-synonyms 
(near isomorphs) of ammotium morenoi acosta. 



Remarks. The specimen illustrated by us in Figs 32-34 was 
not figured or described by Millett but comes from a slide in 
the Millett Collection labelled Haplophragmium cassis Parker 
(BMNH no. 1955.11.1.1118-1133) and was undoubtedly part 
of his concept of that species. It is a typical representative of 
Ammotium morenoi. The small test is axially compressed and 
consists of a short, completely coiled planispiral initial stage, 
followed by an uncoiled portion of about 5 low, elongate, 
laterally compressed chambers which on the interior side 
reach back toward the initial planispire. The single aperture is 
a narrow elongate slit with rounded extremities, situated at 
the apex of the final chamber, in a marginal or outer position. 

Under the name of H. cassis, Millett (1899, pi. 5, figs 4a, b, 
5a, b) did illustrate two specimens, which belong to different 
species of Ammotium. The latter (reproduced here as Fig. 
1.5a,b) is the upper part of an Ammotium pseudocassis 
(Cushman & Bronnimann, 19486)(see p. , below) but was not 
found in the Millett Collection. The former (Fig. 1.4a,b) is a 
complete specimen of A. directum (Cushman & Bronnimann, 
19486) and is refigured in Fig. 31. 

In addition to this specimen, we have also illustrated in Fig. 
54, for the purpose of comparison, the lateral view of a 
typical specimen of A. morenoi from the mangrove sediments 
of Guaratiba, Brazil (see Zaninetti et al. , 1977). It consists of 
an initial, almost involute planispire, followed by two unise- 
rial, laterally flattened, low and elongate chambers, which on 
the inner side extend backwards toward the early spire. 

In common with other brackish foraminifera, A. morenoi is 
highly variable in its overall morphology, in particular in size 
and in outline of the test in lateral view. From small, almost 
triangular forms, as represented by the holotype of morenoi 
or the holotype of Ammoscalaria fluvialis Parker (19526), we 
find all possible transitions to the elongate slender specimens 
of Ammobaculites salsus described by Cushman & Bronni- 
mann (1948a) from Trinidad, or to the large and elongate 
individuals recorded by Poag (1978) from Gulf Coast estuar- 
ies. Brodniewicz (1965: 187-194, text-figs 21-25) has shown 
that a similar form, identified by her as Ammotium cassis 
(Parker), from the Baltic, is also characterized by a great 



1. Ammobaculites salsus Cushman & Bronnimann, 
1948a and A. distinctus Cushman & Bronnimann, 
19486. 

Haman (1983) was the first author to place Ammobaculites 
(=Ammotium) salsus into synonymy with Acosta's species. 
In the introduction to his paper, Acosta (1940: 269) wrote 
that the agglutinating species were rare in the shallow water 
assemblages from the Gulf of Santa Maria, Camaguey Prov- 
ince, Cuba, which were dominated by miliolids and nonion- 
ids. The Gulf of Santa Maria is bordered by extensive 
mangrove swamps. It is therefore assumed that the tests of 
the brackish agglutinated species, such as A. morenoi, had 
been transported by wave action into the marine environment 
of the open Gulf and were not in situ at the locality where 
Acosta collected them. Acosta (1940: 275) claimed to have 
deposited the types of his species in the Cushman Collection, 
which were later transferred from Sharon, Massachusetts to 
the U.S. National Museum of Natural History, Washington, 
D.C. A search by P.B. for the type specimen of A. morenoi 
proved unsuccessful and it seems that Acosta never did 
deposit his types in the Cushman Collection. Acosta's draw- 
ings (op.cit. pi. 49, figs 3,8, non fig. 1) leave no doubt, 
however, that Ammotium morenoi and A. salsum, originally 
described from Trinidad mangrove swamps, are one and the 
same. 

When comparing the two 'species', the apertural view of 
the holotype of Ammotium morenoi (Acosta, 1940: pi. 49, 
fig. 8) is of interest. It shows a slit-like opening at the apex of 
the final chamber, in a marginal or outer position; the same 
type of aperture occurs in A. salsum. In both holotypes the 
peripheral outline of the initial planispire, as seen laterally, is 
perfectly rounded and not angular as in Ammotium distinc- 
tum (Cushman & Bronnimann (19486: 40, pi. 7, fig. 14), 
which has also been described from the brackish mangrove 
sediments of Trinidad. This latter form was originally intro- 
duced as a variety of Ammobaculites salsus. As there are no 
intermediates between distinction and salsum, the former is 
here elevated to specific rank. Authors, however, normally 



114 



P.BRONNIMANN AND J.E. WHITTAKER 



make no distinction between the two (see Phleger, 1954: pi. 
Mig-8). 

We have illustrated in Fig. 55 a lateral view in oil of 
Ammotium distinctum, from the mangrove sediments of 
Acupe, Brazil. The angular outline of the early planispire is 
clearly shown. The test begins with a relatively large prolocu- 
lus of 65 [Am diameter, followed by a larger deuteroloculus of 
75 [xm diameter. The total number of chambers in this 
specimen is eight, including the embryonic chambers. The 
height of the test is 220 \im, width (length) 125 \im, and 
length of aperture 45 |xm. Apart from the distinct angular 
periphery, there are no other important differences between 
Ammotium morenoi and A. distinctum. 

2. Ammoscalaria fluvialis Parker, 19526. 

Parker (1952b: 444, pi. 1, fig. 24) first described this species 
from the Housatonic River, Long Island Sound, depth 3 m. 
From its association with other brackish species in her Facies 
1, such as Trochammina inflata, Jadammina macrescens and 
Miliammina fusca , it can be inferred that A. fluvialis is also a 
brackish-water form. The morphology of the holotype is 
virtually identical with the holotype of A. morenoi, and for 
this reason we regard it as a junior synonym of the latter. 

3. Lituola cassis Parker, in Dawson, 1870. 

We have compared Ammotium morenoi with Lituola cassis 
Parker, the type species of Ammotium Loeblich & Tappan 
(1953). The lectotype of Ammotium cassis (BMNH no. ZF 
4637), designated by Hodgkinson (1992), on our advice, is 
re-illustrated in Figs 38^41. It is from Gaspe Bay, Gulf of St. 
Lawrence, Canada, and came from the W.K. Parker Collec- 
tion; it was collected in 16 fathoms (30 m), which suggests a 
marine environment, but the specimens could have been 
washed in from a brackish locality. Loeblich & Tappan (1987, 
pi. 60, figs 1,2) illustrate a 'Holocene' specimen from off 
Alaska in 223 m of water; should this specimen have been in 
situ it would further undermine the supposedly exclusively 
brackish nature of the genus, a factor that needs further 
investigation. 

The lectotype clearly shows the initial planispire, then the 
uniserial inward slanting narrow and low chambers; the 
oblong aperture is at the apex of the final chamber, in a 
marginal or outer position (see also Goes, 1894, pi. 5, figs 
152-157). The lectotype and paralectotypes are five times 
larger and much more massive than A. morenoi, though the 
two in several other respects are quite similar. It is our 
opinion that A. cassis should only be used for large and 
massive individuals, but at the same time we have our 
reservations that ecological factors (?marine salinities) may 
be responsible for the massive development of the cassis test 
(see also remarks above, on A. cassis sensu Brodniewicz 
(1965) from the Baltic). It is even three times the size of 
Poag's (1978) material from the Gulf Coast estuaries, the 
largest known specimens of A. morenoi from the tropics, 



moreover Poag's specimens are very elongate and com- 
pressed with the uniserial portion quite unlike that of the true 
cassis. 

The dimensions of the lectotype are: maximum height — 
1600 |xm; maximum width — 785 urn; maximum thickness — 
360 |i,m; thickness of planispiral portion — 125 \im. 



4. Ammobaculites prostomum Hofker, 1932. 

This species was described by Hofker (1932: 87-91, text-figs 
14a-f, 15a-d) from the Ammontatura, a part of the Gulf of 
Naples, with a depth of 150-200 m. The shapes of the 
illustrated specimens, seen laterally, particularly the short 
individuals (text-figs 14a and f), much resemble the small 
specimens of Ammotium morenoi such as our Fig. 54. On 
Hofker's short specimens the sutures are not shown fully, but 
on the larger specimens (text-fig. 15e) they are, toward the 
outer margin, at first outward slanting (not inward), then 
parallel up to the end of the uniserial portion. In lateral 
outline, these short specimens are near isomorphs of A. 
morenoi. However, the aperture is not placed asymmetri- 
cally, at the outer margin of the test as in Ammotium, but 
symmetrically in respect to the shape of the final chamber. 
For these reasons, Hofker's species does not belong to 
Ammotium. It is also a marine species and much resembles 
the group pf forms described and illustrated by Hoglund 
(1947, pi. 31, figs la-g) from Bjorkholmen, Gullmar Fjord, 
from a depth of 30 m, under the name of Ammoscalaria 
pseudospiralis (Williamson). 



5. Ammoscalaria pseudospiralis sensu Hoglund, 1947. 

The genus Ammoscalaria was erected by Hoglund (1947: 
151-153) with Haplophragmium tenuimargo Brady (1884) as 
type species. Into his new genus he also placed Proteonina 
pseudospiralis Williamson, 1858. However, Ammoscalaria 
pseudospiralis was described by Hoglund (1947: 159-162, pi. 
31, figs la-p) exclusively from material obtained in the 
Gullmar Fjord, where it occurs commonly, and from the 
Skagerak, not on the basis of Williamson's material which 
was not available to him. The chambers of the rectilinear 
portion of this marine species are 'irregularly rectangular in 
lateral view' and there are 'no external sutures'. The oblong 
aperture is in a symmetrical position in respect to the final 
chamber and not asymmetric, as in Ammotium. We therefore 
do not regard Hoglund's species as a synonym of pseudospira- 
lis, although certain smaller specimens could be regarded as 
isomorphs of Williamson's taxon, particularly when seen in 
lateral view (e.g. pi. 31, figs lm,n). Rather, Hoglund's form 
is most probably a junior synonym of Ammobaculites 
(—Ammoscalaria) prostomum Hofker, 1932. 



Figs 16-21 'Haplophragmoides' wilberti Anderson. Figs 16-18, Side.edge and view of other side (X115). BMNH no. 1911.11.1.5003; Figs 
19-21, Side, edge and view of other side (X160), respectively. BMNH no. ZF 5002. Specimen from Bronnimann sample BR146, Acupe, 
Brazil, for comparison. 

Figs 22-24 Trochammina? milletti sp.nov. Figs 22,23, Detail of aperture (XI, 700) and side view (X320), respectively. Holotype, BMNH no. 
1911.11.1.1088; Fig. 24, Side view (X260). Paratype, BMNH no. 1955.11.1.1089. 

Figs 25-27 Paratrochammina simplissima (Cushman & McCulloch). Spiral, edge and umbilical views (X170). BMNH no. 1955.11.1.1141. 

All from the Millett Collection, Malay Archipelago, except where stated. 



TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 



115 




116 



P.BRONNIMANN AND J.E. WHITTAKER 



Ammotium pseudocassis (Cushman & Bronnimann, 
1948) Figs 1.5, 50, 53 

1899 Haplophragmium cassis (Parker); Millett: 359 (pars); 
pi. 5, figs 5a, b only (non Lituola cassis Parker, 1870). 

19486 Ammobaculites pseudocassis Cushman & Bronni- 
mann: 39, 40; pi. 7, figs 12a,b. 

1983 Ammoscalaria pseudospiralis (Williamson); Haman: 
72; pi. 5, fig. 5 (non Proteonina pseudospiralis Will- 
iamson, 1858). 

Remarks. This species was illustrated by Millett (1899, pi. 5, 
figs 5a, b; here reproduced as Fig. 1.5a,b) under the name of 
Haplophragmium cassis (Parker). It is an upper fragment of 
an elongate test consisting of three rounded (in transverse 
section), hardly compressed chambers. Millett's drawing 
shows the inward and backward trending sutures and the 
rounded aperture in marginal position. Within the concept of 
this species, Millett also included specimens of Ammotium 
directum (Cushman & Bronnimann)(Figs 1.4a,b, 31,?36) and 
Ammoastuta salsa Cushman & Bronnimann (Figs 1.6a,b, 35). 
As already mentioned, the fragment of A. pseudocassis 
illustrated by Millett in his pi. 5, figs 5a, b (see our Figs 
1.5a,b) could not be found in his Malay collection. 

For comparison, we have illustrated typical specimens in 
lateral view of A. pseudocassis from the mangrove sediments 
of Guaratiba, Brazil (see Zaninetti et al. , 1977), one by SEM 
(Fig. 50), the other by optical photography in immersion 
(Fig. 53). In the latter specimen the early spiral is reduced to 
two chambers, a relatively large proloculus of 50 (xm diam- 
eter, and a larger deuteroloculus of about 75 urn diameter. 
The embryonic chambers are not enclosed by other spiral 
chambers, as in A. pseudospirale (Williamson, 1858). The 
total number of chambers, including embryonics, is eight. 

Ammotium pseudocassis differs from A. pseudospirale by 
the elongate, somewhat incurved test, the less compressed 
and elongate chambers and the reduced initial spire. The final 
chamber is usually the dominant one, making up about 
one-third of the test. It narrows toward the oblong aperture 
and extends on the inner side of the test toward the initial 
spiral. The early coil, represented by a reduced spire, consists 
of very few chambers only. A typical embryo consists of two 
very thin-walled chambers, a large proloculus, about 
40-60 um in diameter, and an equally large deuteroloculus. 
The embryo may form all the initial portion of the test. We 
have never found a microspheric specimen of A. pseudocassis 
and where the taxon is frequent, A. pseudospirale is usually 
absent. The aperture of the holotype of A. pseudocassis, 
deposited in the U.S. National Museum of Natural History 
(registration no. 56764), is not as circular as that shown in 
Millett's drawing, but distinctly oblong. As in Ammotium 
cassis and A. morenoi, the aperture is situated at the apex of 
the last chamber in a marginal position (see Cushman & 



Bronnimann, 19486, pi. 7, fig. 12b). However, it seems that, 
when the final chamber is hardly compressed, the aperture 
may become rather centred and more rounded than slit-like, 
but never completely circular. 

Dimensions. Fig. 50: Height of test — 480 \im; maximum 
width — 150 um; height of final chamber — 290 pm; maxi- 
mum diameter of aperture — 50 um. 

Fig. 52: Height of test — 575 pm; maximum width — 170 
um; height of final chamber — 375 mm; maximum diameter 
of aperture — 75 um; thickness of wall (final chamber) 
10 um. 

Environment. The group of forms referred by Millett to H. 
cassis (Parker) occur only at Station 9, Area 1, . . . 'where 
they are not uncommon'. They are all exclusively brackish 
water species. 

Ammotium directum (Cushman & Bronnimann, 1948b) 

Figs 1.4,(?1.7),31,36,37,45-47 

1899 Haplophragmium cassis (Parker): 359 (pars); pi. 5, 
figs 4,?7 only (non Lituola cassis Parker, 1870). 

19486 Ammobaculites directus Cushman & Bronnimann: 
38; pi. 7, figs 3,4. 

1956 Ammotium sp. D. Warren: 139; pi. 1, figs 19-21. 

1957 Ammobaculites directus Cushman & Bronnimann; 
Todd & Bronnimann: 23; pi. 2, fig. 6 only (non Fig. 
7). 

1988 Ammotium casamancensis (sic) Debenay: 46,47; pi. 
1, figs 1-3. 

Remarks. Under the name of H. cassis (Parker), Millett 
(1899, pi. 5, figs 4a, b; here reproduced as Figs 1.4a,b) 
illustrated a specimen of Ammotium directum (Cushman & 
Bronnimann). Our identification is based on the overall 
outline, the shape of the sutures, and the strong lateral 
compression of the test. The specimen illustrated by SEM in 
Fig. 31 is that very same specimen, viewed from the other 
side. 

The fragment shown in Millett's fig. 7, which he compared 
to ... 'a species of Reophax, with the plan of growth and 
chevron-shaped chambers of Frondicularia may be that 
illustrated by SEM in Figs 36,45,46, although the chevron- 
shaped chambers are exaggerated, as they are in the drawing 
of fig. 4a (compare with our Fig. 31). The aperture in both 
specimens is slit-like, without an everted border, and is 
situated at the apex of the final chamber. For comparative 
purposes, a lateral view of a specimen of A. directum, from 
the mangrove sediments of Acupe, Brazil (BMNH no. ZF 
4999) is illustrated in Fig. 47; the asymmetrical sutures are 
well exhibited. 

The tests of Ammotium directum in the Millett Collection 
are extremely fragile, in contrast to those found in Trinidad 



Figs 28-30 Trematophragmoides bruneiensis Bronnimann & Keij. Side, edge and view of other side (X115). BMNH no. 1955.11.1.1136. 
Figs 31, 36, 37 Ammotium directum (Cushman & Bronnimann). Side views of three separate specimens (X 185, 205 and 185, respectively). 

BMNHnos. 1955.11.1.1118-1120. 
Figs 32-34 Ammotium morenoi (Acosta). Side, edge and oblique apertural views (X250). BMNH no. 1955.11.1.1122. 
Fig. 35 Ammoastuta salsa Cushman & Bronnimann. Side view (X 200). BMNH no. 1955.11.1.1121. 
Figs 38-41 Ammotium cassis (Parker). Apertural, oblique apertural, side and edge views (X45). Lectotype, BMNH no. ZF 4637, Gaspe 

Bay, Gulf of St. Lawrence, Canada. 
Figs 42-44 'Ammobaculites' exiguus Cushman & Bronnimann. Fig. 42, Side view (X185). BMNH no. 1955.11.1.1058; Figs 43,44, 

Oblique-apertural and edge views (X175). BMNH no. 1955.11.1.1057. 
All from Millett Collection, Malay Archipelago, except where stated. 



TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 



117 




118 

or Brazil. Cushman & Bronnimann (19486) distinguished two 
species of Ammotium with strong lateral compression, 
namely A. directum and A. diversum. To these has to be 
added a third, A. subdirectum Warren, 1956. 

Ammotium directum, the more common species, as 
described above, has strongly incurved sutures of an asym- 
metrical type with a shorter outer or marginal branch and a 
longer, inner branch, which slants toward the initial spire (see 
Fig. 47). Occasionally, some sort of chevron pattern is 
formed but rarely to the extremes indicated by Milieu's 
drawing (pi. 5, fig. 4a; our Fig. 1.4a). A. directum is always 
characterized by this asymmetrical type of suture. The test, 
moreover, is strongly compressed laterally and the width of 
the flattened chambers does not increase much in the course 
of growth. The aperture is slit-like and situated at the apex of 
the final chamber, more or less in a marginal position. The 
initial spiral consists of several chambers. 

A. diversum is less common than A. directum. The only 
significant difference lies in the sutural shape, which in the 
former in the final ontogenetic stage, is always more or less 
horizontal, slightly incurved and in extremes, no longer 
asymmetrical (see Cushman & Bronnimann, 19486, pi. 7, figs 
5,6). Furthermore, in this species, the sutures of the early 
uniserial portion are slanting inward toward the initial coil. 
Occasionally, there is a suggestion that the two are linked by 
transitional forms. Should the two be ultimately considered 
synonymous then we would prefer to retain A. directum, as 
this, although printed on the same page, was described first. 
For the time being, however, both are retained. 

A. subdirectum was described by Warren (1957, pi. 4, figs 
6-8) from the marshes of the Buras-Scofield bayou region of 
southeastern Louisiana. We have encountered it but rarely in 
the mangrove sediments of Acupe, Brazil and Warren him- 
self (1957: 33) mentioned that . . . 'specimens were rare 
wherever found except in one of the polyhaline marsh 
samples'. Two specimens from Acupe (BMNH nos ZF 5000 
and 5001) are illustrated in Figs 48,49,51,52 for comparison 
with the Malay species of Ammotium. A. subdirectum is a 
many-chambered species characterized by the same asym- 
metrical type of sutures as found in A. directum. In the final 
growth stages the sutures may become more or less symmetri- 
cal and arranged in a chevron-like pattern, as shown in 
Millett's drawing of the fragment illustrated in pi. 5, figs 7a, b; 
Fig. 1.7a,b. Figs 36,45,46 could represent this specimen 
which is part of either an A. subdirectum, or an A. directum 
as discussed above. Normally, A. subdirectum is about twice 
as long as A. directum and composed of more chambers. The 
test is slightly incurved and the width of the chambers, seen 
laterally, increases quite strongly towards the final chamber. 
The aperture is a narrow oblong slit, as in A. directum and A. 
diversum, and situated at the apex of the final chamber, more 
of less in a marginal position (see Figs 48,49,51,52). In all 
three species, the agglutinant is fine-grained and the surface 
of the test usually appears smooth, occasionally even some- 
what glossy. 

Dimensions of figured specimens (BMNH no. 
1955.11.1.1118). Height of test — 290 ^m; maximum width of 
final chamber — 73 \am. 

(BMNH no. 1955.11.1.1119). Height of fragment — 
270 \im; length of apertural slit — 45 \un. 

(BMNH no. 1955.11.1.1120). Height of test — 330 \im. 

Environment. This species was found only at Station 9 



P. BRONNIMANN AND J.E. WHITTAKER 
(Area 1). It is a typical brackish-water species. 

Genus ARENOPARRELLA Andersen, 1951a 

Type SPECIES. Trochammina inflata (Montagu) var. mexi- 
cana Kornfeld, 1931. 

Arenoparrella mexicana (Kornfeld, 1931) 

Figs 1.3,9-12 

1899 Haplophragmium agglutinans d'Orbigny var. triper- 
forata Millett: 358 (pars); pi. 5, figs 3a, b only; non 
figs 2a, b. 

1931 Trochammina inflata (Montagu) var. mexicana Korn- 
feld: 86; pi. 13, figs 5a-c. 

1951a Arenoparrella mexicana (Kornfeld); Andersen: 31; 
fig. la-c. 

19516 Arenoparrella mexicana (Kornfeld); Andersen; 96; 
pi. 11, figs 4a-c. 

1977 Arenoparrella mexicana (Kornfeld); Zaninetti et al.; 
pi. 2, figs 3,7. 

Remarks. One of Millett's illustrated specimens (1899, pi. 5, 
figs 3a, b; reproduced here as Figs 1.3a,b) of H. agglutinans 
var. triperforata is, in fact, a typical specimen of Arenopar- 
rella mexicana (Kornfeld). It is re figured here by SEM (Figs 
9-12) and shows that the original drawing of the edge view in 
particular, is very misleading. As discussed above under the 
description of the lectotype of Acupeina triperforata (Millett), 
the edge view of fig. 3b also suggests that there are only three 
large everted apertures. The reality is an aperture consisting 
of a vertical slit lined by slightly uplifted borders, in an 
interiomarginal position, of about 50 ^m length and 8 \im 
width, and 12 small, irregularly arranged, rounded pores 
above this primary aperture, of between 5 and 10 urn diam- 
eter, devoid of everted rims. Millett's specimen (BMNH no. 
1955.1.1.1075) is tilted so far forward in apertural view that 
the primary vertical slit, so clearly visible in fig. 3a, might not 
have been seen, but it is puzzling to understand why he 
illustrated the apertural pores as he did. Millett's material 
from stations 5 and 9 quite clearly represents both Acupeina 
triperforata and Arenoparrella mexicana, which is not surpris- 
ing as they commonly occur together. The illustrated speci- 
men of the latter is completely involute (hence the small axial 
depression is closed). The final whorl consists of 4, axially 
compressed chambers which gradually increase in size with 
growth. In edge view the periphery is rounded. Umbilical and 
spiral sutures are poorly defined and the agglutination is 
rather fine-grained and produces a smooth surface. 

Dimensions of figured specimen (BMNH no. 
1955.11.1.1075). Maximum diameter — 290 fim; minimum 
diameter — 240 |xm; axial height (thickness) — 120 \im. 

Environment. See under Acupeina triperforata (p. ). 
Arenoparella mexicana is a typical tropical and subtropical 
mangrove swamp species. 



Genus HAPLOPHRAGMOIDES Cushman, 1910 

Type SPECIES. Nonionina canariensis d'Orbigny, 1839. 

Remarks. The wall structure of the type species is unknown; 
we are not even sure of the apertural position, for that 
matter. Although the wall of H. wilberti, the species in the 



TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 



119 



Millett Collection, is of the Trochamminina-type (see below), 
we prefer, in our treatment of this species, to use Haplo- 
phragmoides in inverted commas, until more is known about 
canadensis. 

'Haplophragmoides' wilberti Andersen, 1953 

Figs 1.12, 16-21 

1899 Trochammina ochracea Williamson; Millett: 363, pi. 

5, figs 12a-c (non Williamson, 1858). 
1953 Haplophragmoides wilberti Andersen: 21, pi. 4, figs 

7a,b. 
1961 Haplophragmoides wilberti Andersen; Todd & Low: 

133; pi. 1, fig. 5. 
1973 Haplophragmoides wilberti Andersen; Haynes: 

27-30, pi. 2, fig. 1; pi. 29, fig. 11; text-figs 5.3-7. 
1977 Haplophragmoides wilberti Andersen; Zaninetti et 

al: pi. 1, figs 12,13. 
1981 Trochammina sp., Cann & de Deckker: 668, pi. 2, 

figs 1-19. 
1983 Haplophragmoides wilberti Andersen; Haman: 71; 

pi. 3, figs 14,15. 

Remarks. On re-examination. Milieu's (1899, pi. 5, figs 
12a-c; reproduced here as Figs 1.12a-c) so-called Trocham- 
mina ochracea proved to be a planispiral 'Haplophrag- 
moides'. It is re-illustrated by SEM in Figs 16-18 and it 
(BMNH no 1955.11.1.5003) clearly shows the same collapse 
features as the original drawings. In addition to this speci- 
men, we have illustrated for comparative purposes (Figs 
19-21), another, somewhat less deformed specimen, from 
Acupe, Brazil (BMNH no. ZF 5002). 

The coiling of Millett's species is planispiral, virtually 
involute, with 7 or 8 chambers in the final whorl. The 
aperture is a single interiomarginal equatorial slit with a 
broad everted border. The intercameral sutures are incurved, 
occasionally sinuous. We are placing it into the widespread 
brackish form, 'H'. wilberti Andersen. Millett must have 
regarded it as a Trochammina because of the incurved test 
seen in edge view. 

Cann & de Deckker (1981, pi. 2, figs 1-19) illustrated from 
ephemeral lakes adjacent to the Coorong Lagoon, South 
Australia, a series of haplophragmoid forms, in part 
deformed, which they called Trochammina sp. They are very 
similar to T. ochracea sensu Millett and we have also placed 
them in '//'. wilberti. 

Collapse features occur often in brackish foraminifera. The 
overall consistency of the agglutinated phase, in particular its 
thickness and cementation, seems to play a role. In the 
deformed Millett material it appears that the agglutinated 
phase is rather weakly developed. In non-deformed speci- 
mens of '//'. wilberti, at our disposal, from brackish localities 
in Nigeria and New Guinea, the wall structure was analysed 
using high-resolution scanning electron microscopy of frac- 
tured tests. It was found that the wall of these specimens is 
made up of the organic phase (represented by thin inner and 
outer sheets and material ('glue') between agglutinated ele- 
ments), and the agglutinated phase. There were no perfora- 
tions nor alveolar pseudopores present. This is the 
characteristic Trochamminina-type wall. In these latter, non- 
deformed specimens, the agglutinated phase appears to be 
stronger, perhaps better cemented, than the Millett material 
from the Malay Archipelago. 

Dimensions of figured specimens (MALAY SPECIMEN, 



BMNH no. 1955.11.1.5003). Maximum diameter — 490 urn; 
axial height (thickness) — 125 \xxn. 

(BRAZILIAN SPECIMEN, BMNH no. ZF 5002). Maxi- 
mum diameter — 340 um. 

Environment. According to Millett (1899; 363) this species 
. . . 'has been observed only at Station 3'. It is a good 
brackish water indicator and occurs in association with Trem- 
atophragmoides bruneiensis at this locality. 



Genus PARATROCHAMMINA Bronnimann, 1979 

Type species. Paratrochammina madeirae Bronnimann, 
1979 



Paratrochammina simplissima (Cushman & McCulloch, 
1948) Figs 1.9,25-27 

1899 Haplophragmium nanum Brady; Millett: 360; pi. 5, 
figs 9a-c {non Brady, 1881). 

1939 Trochammina pacifica Cushman var. simplex Cush- 
man & McCulloch: 104; pi. 11, fig. 4 (non Friedburg, 
1902). 

1948 Trochammina pacifica Cushman var. simplissima 
Cushman & McCulloch: 76 (nomen novum). 

1956 Trochammina simplissima Cushman & McCulloch; 
Bandy: 198; pi. 29, figs 14a-c. 

1979 Paratrochammina simplissima (Cushman & McCul- 
loch); Bronnimann: 10; figs 2,3,6A-J,8A-H (q.v. for 
full synonymy). 

Remarks. Millett's illustrated specimen, attributed to H. 
nanum Brady (op.cit. pi. 5, figs 9a-c; reproduced here as Figs 
1.9a-c), is a sinistrally coiled specimen with 5 chambers in the 
final whorl. From the drawings it can be seen that the 
chambers of the final whorl are strongly compressed in an 
axial direction and the ultimate chamber is radially elongate. 
The intercameral sutures are well defined and the agglutinant 
of the spiral side appears to be distinctly coarser than that of 
the umbilical side. The aperture, which is an essential generic 
criterion, is only visible in edge view and its umbilical 
extension, if any, cannot be seen in the drawing of the 
umbilical side. We have searched the Millett Collection to 
find this figured specimen but the closest to it is a dextrally 
coiled individual (Figs 25-27), so it is possible that Millett's 
drawings could be reversed. Our figured specimen is 
undoubtedly Paratrochammina simplissima (Cushman & 
McCulloch). The single umbilical aperture is an elongate 
interiomarginal slit in the final septum, which extends from 
the surface of the first chamber of the final whorl onto that of 
the penultimate chamber. Its length is about 120 yun and it is 
lined by a weakly uplifted border of agglutinated fragments. 
The final whorl has 5 chambers, as in the original drawing, 
but the ultimate chamber, perhaps, is radially not as elongate 
as in Millett's figure. The test consists of 10 chambers, the 
coiling is rather tight and the axial depression (umbilicus) is 
therefore virtually closed. The radial sutures are well defined 
on both sides and the outline of the test is weakly lobate; the 
periphery, as seen in edge view, being compressed but still 
rounded. The spiral side is almost flat and the umbilical side 
slightly concave. As in Millett's illustrated specimen, ours is 
also more coarsely agglutinated on the spiral side than 
umbilically. 
The marine, shallow water P. simplissima differs in all 



120 



P.BRONNIMANN AND J.E. WHITTAKER 



pertinent features (size, chamber inflation and shape, aper- 
ture, etc.) from Brady's deep water species Haplophragmium 
(=Trochammina) nanum which was lectotypified, 
redescribed and illustrated by Bronnimann & Whittaker 
(1980: 177, figs 1-9). P. simplissima is highly variable in the 
overall shape and outline of the test (see Bronnimann, 1979: 
14, figs 6A-J), however it is usually less compressed axially 
than Millett's specimens. 

Dimensions of figured specimen (BMNH no. 
1955.11.1.1141). Maximum diameter — 370 um; minimum 
diameter — 280 um; axial height (thickness) — 90 um. 

Environment. According to Millett (1899: 360), this species 
... 'is most abundant in Area 1'. It is a marginal marine 
species and significantly, was not listed where true brackish 
species such as Acupeina triperforata, Ammoastuta salsa, 
Arenoparrella mexicana, etc. were recorded. 



Genus TREMATOPHRAGMOIDES Bronnimann & Keij, 
1986 

Type species. Trematophragmoid.es bruneiensis Bronnimann 
&Keij, 1986. 

Remarks. The genera Haplophragmoid.es, Cribrostomoides, 
and Discammina are all superficially similar to Tremato- 
phragmoides. Trematophragmoides Bronnimann & Keij is 
slightly evolute and planispiral with 3 apertures per chamber: 
a single primary equatorial interiomarginal aperture and one 
on each side of the chamber, umbilically situated on the 
suture and posteriorly directed. Haplophragmoides Cushman 
(1910) is also planispiral but has only one aperture per 
chamber. Cribrostomoides Cushman (1910) is usually invo- 
lute, with streptospiral coiling initially, becoming planispiral 
in the adult whorls; the aperture is a equatorial, single areal 
slit (with lip) near the base of the septal face, becoming 
subdivided into a linear series of openings in gerontic forms. 
Discammina Lacroix (1932) is planispiral and slightly evolute, 
has a low interiomarginal equatorial aperture and is said to 
have an . . . interior divided by thin straight organic parti- 
tions, not corresponding to the original apertural face and not 
always reflected at the surface' (Loeblich & Tappan, 1987: 
68). 

Trematophragmoides bruneiensis Bronnimann & Keij , 
1986 Figs 1.8, 28-30 

1899 Haplophragmium compressum Goes; Millett: 359; pi. 

5, figs 8a-c {non Lituolina irregularis var. compressa 

Goes, 1882). 
1986 Trematophragmoides bruneiensis Bronnimann & 



Keij: 16; pi. 1, fig. 1-10, pi. 2, figs 3-5, pi. 10, figs 
1-3, text-fig. 1. 

Remarks. Bronnimann & Keij (1986) described from brack- 
ish waters of Brunei, NW Borneo, a planispiral agglutinated 
foraminifer with an interiomarginal and equatorial primary 
aperture and two secondary lateral apertures per chamber. 
The test shape is quite variable and the periphery, as seen in 
edge view, can be broadly rounded or compressed, almost 
subcarinate. Millett's roughly agglutinated species (1899, pi. 
5, figs 8a-c; reproduced here as Fig. 1.8a-c), attributed by 
him to Goes' species H. compressum, shows two lateral 
openings and a single equatorial primary opening per cham- 
ber, and is undoubtedly a Trematophragmoides. Our SEM 
illustrations (Figs 28-30) show the same specimen as that 
drawn by Millett. As the early radial sutures are indistinct, 
the total number of chambers cannot be determined with 
certainty; the final whorl, however, contains 6 chambers. The 
lateral secondary apertures are well exhibited in Fig. 30 and 
the equatorial primary aperture in Fig. 29. Although the 
number of chambers in the final whorl is less than in the types 
of T. bruneiensis, other features agree well and there is no 
doubt that the two are one and the same. 

Dimensions of figured specimen (BMNH no. 
1955.11.1.1136). Maximum diameter — 470 um; maximum 
thickness (final chamber) — 200 pm. 

Environment. This species occurs only at Station 3. Millett 
(1899) does not offer any information about its association 
with other species, but from a study of his collection it can be 
seen to occur with 'Haplophragmoides' wilberti. From this 
information, Station 3 must have been a brackish locality. 



Genus TROCHAMMINA Parker & Jones, 1859 
Type species. Nautilus inflatus Montagu, 1808. 



Trochammina? milletti sp.nov. 



Figs 1.10,22-24 



1899 Haplophragmium anceps Brady; Millett: 361, pi. 5, 
figs 10a,b {non Brady, 1884). 

Diagnosis. Small conical, tightly-coiled Trochammina? with 
three large subglobular chambers in the final whorl. 

Name. In honour of Fortescue William Millett. 

Holotype. BMNH no. 1955.11.1.1088. Illustrated in Figs 
22,23. This may be the specimen figured by Millett (1899, pi. 
5, figs 10a, b; reproduced here as Figs l.lOa.b). From Station 
12, Area 1. 

Description (holotypes). Test free, dextrally coiled coni- 
cal trochospire with pointed initial portion. Final volution 



Figs 45-47 Ammotium directum (Cushman & Bronnimann). Figs 45,46, Detail of aperture (X975 and X280, respectively). BMNH no. 

1955.11.1.1119; Fig. 47, Side view (X160). BMNH no. ZF4999, mangrove sediments, Bronnimann sample 93, Acupe, Brazil 
Figs 48, 49, 51, 52 Ammotium subdirectum Warren. Figs 48,49, Side and edge views (X85). BMNH no. ZF 5000; Figs 51,52, Side and 

oblique-apertural views (X85 and 125, respectively). BMNH no. ZF 5001. Both from mangrove sediments, Bronnimann sample 93, Acupe, 

Brazil. 
Figs 50, 53 Ammotium pseudocassis (Cushman & Bronnimann). Side view (X160) and separate specimen in clearing oil (X265). 

Bronnimann Collection, mangrove sediments, Guaratiba, Brazil. 
Fig. 54 Ammotium morenoi (Acosta). Side view (X205). Bronnimann Collection, mangrove sediments, Guaratiba, Brazil. 
Fig. 55 Ammotium distinctum (Cushman & Bronnimann). Side view in clearing oil (X330). Bronnimann Collection, sample 145, from 

Acupe, Brazil. 
All from Millett Collection, Malay Archipelago, except where stated. 



TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 



121 




122 



P. BRONNIMANN AND J.E. WHITTAKER 



triserial, consisting of large subglobular chambers, somewhat 
compressed in axial direction, making up major part of the 
test. Coiling tight and axial depression (umbilicus) closed. 
Sutures well defined. Single interiomarginal aperture a small 
arch resting with its slightly upturned border completely on 
surface of first chamber of final whorl. Agglutinant rather 
coarse. 

Dimensions (holotype). Height of test — 160 urn; width 
(umbilical diameter) — 150 |i,m. 

Remarks. Millett (op.cit., pi. 5,, figs 10a, b) attributed this 
small, rather fragile, conical form, to Haplophragmoides 
anceps Brady. The aperture is a broadly rounded interiomar- 
ginal arch, sitting with its border completely on the final 
whorl and therefore the species should belong to Trocham- 
mina (see Bronnimann et al., 1983). Fig. 23 illustrates a 
typical specimen from the Millett Collection but there are 
extremes (Paratype BMNH no. 1955.11.1.1088; Fig. 24) 
where the height of the trochospire and the umbilical diam- 
eter are about the same or the former even appears to be 
slightly larger. In 1983, Bronnimann et al. held great store by 
the fact that in the Trochamminacea the umbilical diameter 
was invariably greater than the length of the axis of coiling 
(height of the trochospire), whereas in the Ataxophragmia- 
cea the reverse was true. This is the first time we have found a 
species, and apparently a single population, at the borderline 
of the two groups. For this reason we have only tentatively 
placed this interesting species in Trochammina. 

The true H. anceps Brady, 1884 is the type species of 
Globotextularia Eimer & Fickert, 1899. This is a robust, deep 
water form, much larger than Millett's species, with a very 
high, often irregular coil, an open umbilicus and larger 
aperture. 

Environment. According to Millett (1899: 361), . . . 'speci- 
mens [of "H. anceps'] are numerous and well distributed'. 
They are found at stations 5,11,12 (Area 1) and 27,28 (Area 
2). It is associated with the agglutinating foraminifera Ammo- 
baculites exiguus at stations 12 and 27 and rare Acupeina 
triperforatai ' Arenoparrella mexicana at Station 5. The former 
is found in both marginal marine and brackish localities, 
whereas the latter are true brackish forms. It is therefore not 
known for certain whether T. ? milletti is a marine or a 
brackish species. 



Genus TRUNCULOCAVUS gen.nov. 

Type species. Trunculocavus durrandi sp.nov. 

Diagnosis. Test free, initially biserial, then abruptly unise- 
rial. Biserial chambers subglobular, uniserial chambers with 
circular transverse section. Wall agglutinated, of 
Trochamminina-type. Aperture single, terminal, circular and 
large, devoid of everted border. 

Name. Derived from the Latin: cavus, a hole or hollow, and 
trunculus, tip or end. 

Remarks. Our new genus has the basic morphology of 
Bigenerina d'Orbigny, 1826 (type species B. nodosaria d'Or- 
bigny, 1826), but differs in the large circular aperture of the 
final chamber of the uniserial stage, devoid of a border 
structure. In contrast, the terminal aperture of B. nodosaria 
is a small central porus with everted border. According to 
Loeblich & Tappan (1987: 172), Bigenerina also has a perfo- 



rate ('canaliculate') wall, whereas Trunculocavus has a 
Trochamminina-type wall. 

In the Millett material, there are well-preserved specimens 
of Trunculocavus durrandi showing an organic structure 
within the large rounded aperture. This structure is different 
from the inner organic sheet (inner organic lining in the sense 
of Bender, 1989: 278), which occurs along the inside of the 
wall of the Trochamminina, because it is independent of the 
agglutinated-organic wall proper. It is suggested that it repre- 
sents the epidermal layer of the protoplasmic body of the 
foraminifer. Therefore, we must distinguish between this type 
of organic structure, as part of the protoplast, and the inner 
organic sheet which covers the inside of the agglutinated wall 
of the Trochamminina-type test (see Bronnimann & Whit- 
taker, 1988), which, although has also been produced by the 
protoplast, is not directly part of it. 

This organic structure, or the epidermal layer of the 
protoplasmic body, occurs inside the terminal aperture of the 
test, either as a large rounded opening limited by a thickened 
border (Fig. 7), or it closes the aperture of the test completely 
and reveals 6 small perforations with tube-like extensions 
(Fig. 1) along the apertural periphery. This organic structure 
does not have a counterpart in the agglutinated-organic phase 
of the wall, another reason for separating it nomenclatorally 
from the inner organic sheet. In fossil specimens, the epider- 
mal layer of the protoplast will naturally be absent, so it could 
not be considered taxonomically. At present, therefore, it has 
no standing in the systematic treatment of these agglutinated 
foraminifera, which is based on test features alone. It should, 
however, be remembered that this situation would have to be 
modified once it becomes possible to take into consideration 
the features of the living organism. 

In a paper by Petrucci et al. (1983: 72-75), there is a 
taxonomic appendix by Medioli, Scott & Petrucci. In this 
appendix a new species, Polysaccammina hyperhalina, is 
introduced which is of interest here because it shows organic 
features similar to those described for T. durrandi. P. hyper- 
halina has a large circular terminal aperture with an irregu- 
larly finished border, devoid of particular border structures. 
Medioli et al. (1983: 72, pi. 21, figs 2,3,6,8) described the 
aperture as invaginated ... 'to form an inner, backward 
pointing funnel'. Their pi. 1, figs la, 2a show the large, 
rounded aperture is closed on the inside, as in T. durrandi, by 
an organic structure having in its centre a small circular porus 
with everted border. Also, as in T. durrandi, this small 
opening appears to be a different from the aperture of the test 
and that it represents the epidermal layer of the protoplast, 
with features which have no counterpart in those of the test 
wall and which is different from the inner organic lining. 



Trunculocavus durrandi sp.nov. 



Figs 2.1,3-8 



1900 Bigenerina digitata d'Orbigny var. Millett: 6, pi. 1, 
figs la,b (non Bigenerina (Gemmulina) digitata d'Or- 
bigny, 1826). 

Diagnosis. As for genus; Trunculocavus is presently mono- 
typic. 

Name. In honour if A. Durrand FRMS, the collector of the 
Malay Archipelago foraminifera described by Millett. 

Holotype. BMNH no. 1955.11.1.187. Illustrated in Figs 
3,4. From Station 9, Area 1. 

Description (holotype. Test free, small and elongate; ini- 



TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 



123 



tially a subglobular protoconch, followed by 4 pairs of 
subglobular, biserial chambers, then abruptly uniserial with 3 
cylindrical chambers. Aperture large, terminal and central 
without everted border; the inner organic sheet closing the 
aperture, however, develops around its circumference, 6 
minute pores with tubular borders. Agglutinated wall of 
granular, but overall smooth appearance. Tubular organic 
pores have no counterpart in agglutinated phase. 

Dimensions (holotype). Height of test — 270 \xm; maxi- 
mum width of test — 75 |im; diameter of aperture — 35 u.m; 
diameters of tubular pores around circumference of aperture 
— 4-5 \im. 

Paratypes. Two paratypes (BMNH nos. 1955.11.1.188,189) 
are illustrated in Figs 5-8. In side view, they are as the 
holotype with a short biserial stage followed by the uniserial 
stage composed of 2 or 3 cylindrical chambers. Paratype 
(BMNH no. 1955.11.1.188; Figs 6-8) shows an aperture 
where the inner organic sheet does not close the opening. The 
sheet itself has an opening, bordered by a thickened rim, 
which is virtually of the same diameter as the rounded 
terminal aperture of the agglutinated phase; there are no 
minute pores as in the holotype. 

Dimensions (paratypes). (BMNH no. 1955.11.1.188) 
Height of test — 230 [xm; maximum width — 90 urn; diam- 
eter of aperture — 40 (am. 

(BMNH no. 1955.11.1.189) Height of test — 240 urn; 
maximum width 75 um. 

Remarks. Millett's actual figured specimen (1900, pl.l, figs 
la,b; reproduced here as Figs 2.1a,b) could not be recognised 
with certainty. Millett's drawing, however, shows a specimen 
with a rather indistinct biserial initial portion of 4 or 5 pairs of 
chambers, then a 4 or 5-chambered uniserial stage; the 
uniserial chambers are cylindrical and the large terminal 
rounded aperture is devoid of an everted border. 

Environment. According to Millett (1900: 6), this species is 
. . . 'confined to Station 9, and the examples, although 
minute, are moderately abundant'. From the same locality 
Millett (1899: 358,359) also found Acupeina triperforata, 
Arenoparrella mexicana, 'Ammobaculites' exiguus, Ammoas- 
tuta salsa and Ammotium spp., all brackish, mangrove 
sediment-dwelling species. It is therefore assumed that T. 
durrandi also lives in a brackish habitat. 



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Bull. nat. Hist. Mus. Lond. (Zool.) 59(2): 125-170 



Issued 25 November 1993 



Foregut anatomy, feeding mechanisms, 
relationships and classification of the 
Conoidea (= Toxoglossa) (Gastropoda) 

JOHN D. TAYLOR 

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5 BD 

YURI I. KANTOR 

A.N. Severtzov Institute of Animal Evolutionary Morphology and Ecology, Russian Academy of 
Sciences, Lenin Avenue 33, Moscow 117071 

ALEXANDER V. SYSOEV 

Institute of Parasitology, Russian Academy of Sciences, Lenin Avenue 33, Moscow 117071 

CONTENTS 



Introduction 125 

Material and Methods 1 26 

Foregut anatomy 126 

Functional morphology of the digestive system and feeding mechanisms in the Conoidea 145 

Relationships of Conoidea 150 

Classification of Conoidea and diagnoses of higher taxa 156 

Acknowledgements 161 

Appendix 1. Description of shell characters used in phylogenctic analysis 162 

Appendix 2. Classification of Recent turrid genera 163 

Refe rences 1 68 



Synopsis. A survey of the anterior alimentary system of species from all the higher taxa of the highly diverse 
gastropod superfamily Conoidea (including the Turridae. Tercbridae, and Conidae) has revealed a great variety of 
foregut structure. A series of anatomical characters of the rhynchodeum, proboscis, buccal mass, radular apparatus 
and foregut glands has been defined and their distribution established amongst the various conoidean families and 
sub-families. Twelve major types of foregut structure were recognised, which ranged from gastropods with a full set 
of foregut organs and glands to others in which most of the structures including the radula, venom gland and 
proboscis are absent. A set of these anatomical characters together with a few shell characters were used in a 
cladistic analysis attempting to determine relationships amongst the conoidean higher taxa. A classification 
incorporating the new anatomical data and based partly upon the phylogenetic analysis recognises 6 families and 13 
subfamilies of Conoidea. New data suggest that the Pervicaciinae and Terebrinae share a common ancestor and 
there is little evidence to justify familial separation of the Coninae. Some major foregut structures seem to have 
evolved independently in different clades. Thus, hollow 'hypodermic' radular teeth have been derived indepen- 
dently in a least five clades; the radular caecum and rhynchodeal introvert have evolved independently in two clades. 
Several clades also show loss of major foregut structures such as the proboscis, venom gland and radular apparatus. 
Finally, the 378 genera of Recent Turridae' are placed into the higher taxa recognised in the proposed classification. 



INTRODUCTION 



The prosobranch gastropod superfamily Conoidea (=Toxo- 
■glossa), which includes the families Turridae, Conidae, Per- 
vicaciidae and Terebridae, is extremely diverse, with as many 
as 679 genera and 10,000 living and fossil species claimed for 
the Turridae alone (Bouchet, 1990) and Conus with around 
500 living species, is considered to be the most diverse genus 
af marine animal (Kohn, 1990). Current classifications of 
axa within the Conoidea are based almost entirely upon shell 



characters, or upon a combination of shell and radular 
characters (Turridae-Powell, 1966; McLean 1971; Kilburn, 
1983, 1985, 1986, 1988; Terebridae-Bratcher & Cerno- 
horsky, 1987). The Turridae are the most morphologically 
disparate of the four families with seventeen subfamilies in 
current use. However, most of these subfamilies are rather 
poorly defined. Despite the biological interest in the venom 
apparatus of the group, little is known of the relationships of 
the Conoidea to other gastropods, of relationships between 
the families of the Conoidea or of relationships within the 
constituent families. 



©The Natural History Museum, 1993 



126 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



The Conoidea are considered to be monophyletic, because 
the families share the common apomorphy of a venom 
apparatus con-sisting of the venom gland and muscular bulb. 
This is thought to have been lost in some taxa, such as some 
highly-derived members of the Daphnellinae and Terebridae 
(Kantor & Sysoev 1989; Taylor, 1990) and all Strictispirinae. 

Compared with the number of living species and the 
attention paid to the description of shells, particularly of 
Conidae, there have been very few anatomical studies of 
Conoidea. However, recently, a much wider range of species 
from the Turridae, Terebridae and Pervicaciidae (Sysoev & 
Kantor 1987, 1988, 1989; Kantor & Sysoev, 1989; Miller, 
1989, 1990; Kantor, 1990; Taylor, 1990) have been investi- 
gated anatomically. These studies illustrate the great variety 
of foregut anatomy, particularly within the Turridae and 
Terebridae. By comparison, the Conidae appear to have a 
relatively uniform foregut anatomy (Marsh, 1971; Miller, 
1989), although they have been surprisingly little studied. 

Until recently, attempts to use anatomical characters in 
determining relationships amongst conoideans were con- 
strained either by the limited range of taxa that had been 
studied or by the small number of characters used. For 
example, an evolutionary scenario for the Conoidea based 
upon characters of foregut anatomy was proposed by Sheri- 
dan etal. (1973), but species were studied from only three out 
of the seventeen turrid subfamilies. Additionally, Shimek & 
Kohn (1981) used only radular characters to produce a 
cladistic analysis of a wider range turrid taxa. 

Another problem in comparing the different taxa studied 
within the Conoidea, is that the nomenclature for the differ- 
ent anatomical structures is inconsistent and very confused. 
This has hampered the recognition of homologous structures 
that may be shared between the different taxa. 

In this paper we attempt a comparative review of the 
anatomy and functional morphology of the conoidean 
foregut. We have attempted to examine species from all the 
currently-recognised subfamilies of Turridae, many species of 
Terebridae, Pervicaciidae and a few species of Conns. Addi- 
tionally, we have incorporated previously published studies 
into our review and attempted to standardize the nomencla- 
ture of the anatomical structures. 

The overall objectives of the study are, firstly, to evalu-ate 
the use of characters of foregut anatomy in determining 
relationships among the Conoidea and secondly, to propose a 
new classification of conoidean higher taxa which incorpo- 
rates these anatomical characters. Foregut anatomy was 
chosen as the focus for this study, because a few previous 
studies (Sheridan et al. 1973; Kantor, 1990) had drawn 
attention to the diversity and complexity of the digestive 
system. As far as is known, other organ systems are similar to 
other neogastropods. 



descriptions in our analysis. Additionally, radular prepara- 
tions were made from a range of other species. 

Critical-point dried preparations for scanning electron 
microscopy were made of some anatomical structures and 
some small species (methods in Taylor & Miller, 1989). 
Radula preparations for both light and scanning microscopy 
were made by standard methods. 



FOREGUT ANATOMY 



A generalized diagram of the conoidean foregut (Fig. 1) 
shows the relative positions of the major structures. 



rstm 



rcoel 




rw 






MATERIAL AND METHODS 

The material on which this study is based consists mainly of 
longitudinal serial sections of the foreguts of a wide range of 
gastropods from all of the currently recognised subfamilies of 
Turridae, many Terebridae and Pervicaciidae and a few 
species of Conidae (Table 1). Dissections were also made of 
most of these species. Also indicated in Table 1 are species 
for which we have used previously published anatomical 



Fig. 1 Composite diagram of the foregut of a hypothetical 

conoidean gastropod showing the location of the major structures 
discussed in the text. No single gastropod possesses all these 
features. Abbreviations: as, anterior sphincter of buccal tube; bl, 
buccal lips; bm, buccal mass; bt, buccal tube; is, intermediate 
sphnicter of buccal tube; m, mouth; mb; muscular bulb; oe, 
oesophagus; p, proboscis; rcoel, rhynchocoel; rs, radular sac; rsp, 
rhynchostomal sphincter; rstm, rhynchostome; rw, rhynchodeal 
wall; s, septum; sg, salivary gland; tm, transverse muscles of 
rhynchodeal wall (shown in part only); vg, venom gland. 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



127 



Table 1. List of species examined in this study. The classification in the list is traditional and follows Powell (1966), McLean ( 1971) and 
Kilburn (1983-89). A new classification is given at the end of this paper. The prefix 'a' denotes species that were studied anatomically and 
the prefix 'r' denotes species for which only the radula was examined. In most cases, animals were both dissected and serial sections made 
of the anterior alimentary systems. 



Pseudomelatominae 

aPseudomelatoma penicillata (Carpenter, 1864). Punta San 

Bartoleme, Mexico. 

aHormospira maculosa (Sowerby, 1834). Sonora, Mexico 

Drilliinae 

aClavus unizonalis (Lamarck, 1822). Lizard I., Queensland, 

Australia. 

aClavus sp. (undescribed species). Guam. 

aSplendrillia chathametisis Sysoev & Kantor, 1989. Chatham Rise, 

South Pacific. 

xDrillia cydia (Bartsch, 1943). British Virgin Islands. 

rDrillia rosacea (Reeve, 1845). West Africa. 

rlmaclava unimacidata (Sowerby, 1834). Baja California, Mexico. 

rSpirotropis monterosatoi (Locard, 1897). East Atlantic. 

xCrassopleura maravignae (Bivona, 1838). Naples, Italy. 

Clavatulinae 

aToxoclionella tumida (Sowerby, 1870). South Africa. 
aClionella sinuuta (Born, 1778). Oudekraal, South Africa. 
aClavatula caerulea (Weinkauff, 1875). Sierra Leone, West Africa. 
aClavatula muricata (Lamarck, 1822). Dakar, Senegal. 

Turrinae 

rLophiotoma acuta (Perry, 1811). Lizard I., Queensland, Australia. 
aGemmula deshayesi (Doumet, 1839). Hong Kong. 
xGemmula kieneri (Doumet, 1840). Hong Kong. 
aLophiotoma leucotropis (Adams & Reeve, 1850). Hong Kong. 
aPolystira albida (Perry, 1811). Caribbean. Data from Leviten 
(1970). 

Cochlespirinae 

aTurricula javana (Linnaeus, 1767). Hong Kong. 
aTurricula nelliae spurius (Hedley, 1922). Hong Kong. 
aAforia abyssalis Sysoev & Kantor, 1987. North-East Pacific. 
aAforia lepta (Watson, 1881). South Pacific, nr New Zealand. 
aAforia inoperculata Sysoev & Kantor, 1988. North-East Pacific. 
alrenosyrinx hypomela (Dall, 1889). East Atlantic. 
aAntiplanes sanctiioannis (Smith, 1875). Okhotsk Sea. 
xAnliplanes vinosa (Dall, 1874). Sakhalin Bay, Okhotsk Sea. 

Crassispirinae 

r Austrodrillia angasi (Crosse, 1863). Sydney, Australia. 
aFuna latisinuata (Smith, 1877). Hong Kong. 
alnquisitor spp. Indian Ocean. 

aVexitomina garrardi (Laseron, 1954). Sydney, Australia. 
xPtychobela griffithi (Gray, 1834). Karachi. 

Strictispirinae 

aStrictispira paxillus (Reeve, 1845). British Virgin Islands. 

xStriclispira stillmani Shasky, 1971. Panama. 

xCleospira ochsneri (Hertlein & Strong, 1849). Galapagos Islands. 

Zonulispirinae 

aPilsbryspira nympha (Pilsbry & Lowe, 1932). Sonora, Mexico. 

Borsoniinae including Mitrolumninae (fide Kilburn, 1986) 

aLovellona airamentosa (Reeve, 1849). Guam. 

aAnarithma metula (Hinds, 1843). Indian Ocean. 

aBorsonia ochraea Thiele, 1925. Indian Ocean, nr Zanzibar 740m. 

aMicanthapex parengonius (Dell, 1956). South Pacific, nr New 

Zealand. 

aTomopleura reevei (C.B. Adams, 1850). Indian Ocean. 

aSuavodrillia kennicotti (Dall, 1871). Japan Sea. 

aTropidoturris anaglypta Kilburn 1986. Southern Indian Ocean. 

aTropidoturris fossata notialis Kilburn, 1986. South Africa. 

aOphiodermella inermis (Hinds, 1843). Bremerton, Washington. 

aOphiodermella ogurana (Yokoyama, 1922). Japan Sea. 



Clathurellinae 

aGlyphostoma Candida (Hinds, 1843). Sonora, Mexico. 

Mangeliinae 

aMangelia brachystoma (Philippi, 1844). Galway, Ireland. 
aMangelia nebula (Montagu, 1803). Galway, Ireland. Also data 
from Sheridan el al. (1973) & Delaunois & Sheridan (1989). 
aMangelia powisiana (Dautzenberg, 1887). Plymouth, England. 
aEucithara stromboides (Reeve, 1846). Guam. 
aHemilienardia malleti (Recluz, 1852). Guam. 
aParamomana cf. rufozonata (Angas, 1877). Rottnest I., Western 
Australia. 

Oenopotinae 

aOenopota levidensis (Dall, 1919). Washington. Data from Shimek 

(1975) 

xPropebela rugulata (Moller, 1866). White Sea. 

Daphnellinae 

aComarmondia gracilis (Montagu. 1803). Brittany, France. Data 

from Sheridan et al. (1973) 

aDuphnella reeveana (Deshayes, 1863). Guam. 

aGymnobela emertoni (Verrill & Smith, 1884). Eastern Atlantic 

Ocean. 

aTereiiopsis levicarinatus Kantor & Sysoev, 1989. Eastern Atlantic 

Ocean. 

aAbyssobela aloxica Kantor & Sysoev, 1986. Northern Pacific 

Ocean. 

aGymnobela latistriaia Kantor & Sysoev, 1986. Northern Pacific 

Ocean. 

aGymnobela oculifera Kantor & Sysoev, 1986. Northern Pacific 

Ocean. 

aPontiothauma abvssicola Smith. 1895. Indian Ocean. Data from 

Pace (1901). 

aPontiothauma mirabile Smith. 1895. Indian Ocean. Data from Pace 

(1901) 

Conorbinae 

aBenthofascis biconica (Hedley, 1903). Sydney, Australia. 
aGenota milraeformis (Woods, 1828). West Africa. 
aGenota nicklesi Knudsen, 1952. West Africa. 

Thatcheriinae 

aThatcheria mirabilis Angas, 1877. North Western Australia. 

Taraniinae 

aTaranis moerchi (Malm, 1861). Sweden. 

Conidae 

aConus flavidus Lamarck, 1810. Queensland, Australia, Data from 

Marsh (1971) 

aConus ventricosus Gmelin, 1791. Tunisia. 

Pervicaciidae 

aPervicacia capensis (Smith, 1873). South Africa. 

aPervicacia kieneri (Deshayes, 1859) Albany, Western Australia. 

aPervicacia tristis (Deshayes, 1859). New Zealand. 

aDuplicaria colorata Bratcher, 1988. Western Australia. 

aDuplicaria duplicata (Linnaeus, 1758). Kenya. 

aDuplicaria spectabilis (Hinds, 1844). Hong Kong. 

a'Terebra' nassoides Hinds, 1844. Oman. 

Terebridae 

aHastula aciculina (Lamarck, 1822). Ghana. 
aHastula bacillus (Deshayes, 1859). Phuket, Thailand. 
aTerebra affinis Gray, 1834. Guam. 
aTerebra babylonia Lamarck, 1822. Guam. 
aTerebra gouldi Deshayes, 1857. Hawaii. 
aTerebra maculata Linnaeus, 1758. Guam. 
aTerebra subulata Linnaeus, 1767. Maldives. 



128 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



Characters of the rhynchocoel 

In all toxoglossans there is a permanent cavity in the anterior 
part of the body called the rhynchodeal cavity or rhynchocoel 
(Fig. 1). It contains the proboscis and is maintained even 
when the proboscis is extended. The rhynchodeal cavity 
opens to the exterior via the rhynchostome, which is situated 
at the ventral margin of the head. The walls of the rhynchoc- 
oel (rhynchodeum) are usually thick and muscular. 

Rhynchostomal sphincter 

This an annular, muscular sphincter which encircles the 
mouth of the rhynchocoel (Fig. 1). It is present in most 
species of Turridae, Terebridae, Pervicaciidae and Conidae, 
but absent in the turrids Clavatula diadema and Tomopleura 
violacea and the pervicaciids Pervicacia tristis, 'Terebra' nas- 
soides, and 'T. ' capen-sis. In these latter pervicaciids and 
some turrids without a prominent sphincter, for example 
Tomopleura, the anterior part of the rhynchodeum is very 
muscular. 




Fig. 2 Ophiodermella inermis; longitudinal section of the anterior 
rhynchodaeum showing the posteriorly-situated, rhynchostomal 
sphincter located on an introvert-like structure. Abbreviations: in, 
introvert; m, mouth; p, proboscis; r, rhynchostome; s, sphincter. 



Position of rhynchostomal sphincter 

In the normal condition, the sphincter is usually situated 
around the rhynchostome, but in some turrids (for example in 
Glyphostoma, Borsonia, Lophiotoma, Pontiothauma and 
Thatcheria) it is situated more posteriorly. In Ophiodermella 
inermis (but not O. ogurana) and Suavodrillia kennicotti the 
moderately large, posteriorly situated, rhynchostomal sphinc- 
ter is probably able to evert, forming a sort of 'rhynchostomal 
introvert' but situated in the middle part of the rhynchocoel 
(Fig. 2). The ability to evert is indicated by the presence of a 
well-developed layer of longitudinal muscles underlying the 
epithelium and by the existence of free space between the 
sphincter and the longitudinal muscle layers. This structure 
may demonstrate the possible origin of the true rhynchodeal 
introvert (see below) or alternatively be an autapomorphy for 
the species. 

Rostrum 

In the some fish-feeding species of Conus, the anterior part of 
the rhynchocoel is elastic and can be greatly extended to 
accomodate large food items during preliminary digestion. 
This extensible feature, known as the rostrum, cannot be 
inverted into the rhynchocoel. 

Rhynchodeal introvert (= labial tube or 
pseudoproboscis) 

In this structure, the rhynchostomal lips are mobile and can 
be retracted into the rhynchocoel by infolding, or extended as 
a tube (Figs 3 & 4). The introvert is found in nearly all the 
species which we and others have studied from the turrid 
sub-family Daphnellinae, e.g. Philbertia linearis, P. leufroyi, 
P. gracilis, Cenodagreutes, Daphnella reeveana (Smith, 1967; 
Sheridan et ai, 1973; unpublished observations), in Hemi- 
lienardia mailed (Mangeliinae) and in all Terebridae and 
Pervicaciidae (Miller, 1975, 1980; Taylor, 1990). We have not 
seen an introvert in any other subfamily of Turridae (except 
perhaps for Ophiodermella, see above), or in the Conidae. In 
species of Daphnellinae the introvert is fairly short, but in 
some terebrids, for example Terebra maculata, the introvert 




Fig. 3 Hemilienardia malleti; extended rhynchodeal introvert, 
forming a pseudoproboscis in a relaxed, critical-point dried 
specimen. Scale bar = 100 um. 

is very long, and when retracted, lies coiled in the rhynchoc- 
oel (Miller, 1970). 

In those animals possessing a rhynchodeal introvert, the 
outer and inner walls are joined by radial muscles (Fig. 5). In 
Turridae, the possession of an introvert is associated with a 
reduction in size or complete loss of the proboscis. However, 
within the Terebridae, even those species with a well- 
developed proboscis possess an introvert. 

Epithelium of the rhynchodeum 

In some Turridae, there is a distinct division in the character 
of the epithelium lining the inner wall of the rhynchocoel. In 
the anterior part of the cavity the epithelial cells are high and 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



129 




0-1 mm 



ovg 



cis, when withdrawn, lies behind the septum, with the 
retracted introvert lying to the anterior. A probably homolo- 
gous septum is also found at the extreme posterior and 
ventral end of the rhynchocoel in Thatcheria and Pontio- 
thauma (Pace, 1901). A thin septum is also found in the 
posterior part of the rhynchocoel in Pervicacia tristis (not 
reported by Rudman (1969)) and in Duplicaria kieneri (Tay- 
lor, unpublished). 

The function of the septum is unknown, but it appears 
better developed in species with a long proboscis and where 
the proboscis withdraws behind the septum. 

Accessory proboscis structure 

This is an extensible muscular structure which arises from the 
left hand wall of the rhynchocoel. It has been found only a 
few species of Terebridae and Pervicaciidae. It is long and 
branched in Hastula bacillus (Taylor & Miller, 1990), shorter 
and club-like in Terebra affinis (Miller, 1971), 'Hastula' 
colorata and D. kieneri and a curved, club-shape in Terebra 
imitatrix (Auffenberg & Lee, 1988). In H. bacillus the 
accessory proboscis is covered in possible chemosensory 
structures (Taylor & Miller (1990). 

Snout gland 

This is a subspherical gland which opens into the right-hand 
posterior end of the rhynchocoel in a number of Conns 
species (Marsh, 1971). The gland consists of folded glandular 
epithelium (Fig. 7) and is surrounded by a muscular sheath of 
circular muscles. From histochemical tests. Marsh (1971) 
concluded that the gland secretes mucus. The gland has been 
reported in 18 species of Conus, all but one of which are 
known to be vermivorous (Marsh, 1971). 



Fig. 4 Daphnella reeveana; A, longitudinal section through the 
foregut; B, Enlargement of the mouth area showing the short 
proboscis lying behind the septum. Abbreviations: bm, buccal 
mass; cm, columellar muscle; con, circum-oral nerve ring; in, 
rhynchodeal introvert; oe, oesophagus; ors, opening of radular 
sac; ovg, opening of venom gland; p, proboscis;s, rhynchostomal 
sphincter; spt, septum; vg, venom gland. 

glandular (Fig. 6C), but in the posterior half the epithelium is 
low, cuticularized and similar in morphology to that of the 
outer surface of the proboscis. This feature indicates that the 
posterior part of the rhynchodeum can be extended outwards 
when the proboscis is protruded through the rhynchostome. 
We have observed this condition of the rhynchocoel epithe- 
lium in Clavatula, and Clionella (Clavatulinae), Vexitomina 
(Crassispirinae), Turricula nelliae spurius (Cochlespirinae), 
Pilsbryspira nympha (Zonulispirinae). and Anarithma metula 
(Borsoniinae). 

In 'lower' turrids, excepting Vexitomina, this feature seems 
to associated with those species in which the buccal mass lies 
in a distal position within the proboscis (see below). Its 
presence may be connected with the mechanism by which the 
buccal mass is everted from the proboscis tip. 

Septum in rhynchodeum 

A thin, slightly muscular septum, pierced by a circular orifice, 
and dividing the rhynchodeal cavity into two parts is known in 
Daphnella reeveana (Fig. 4), Philbertia purpurea (Sheridan et 
al., 1973) and Terebra subulata (Taylor, 1990). The probos- 



The proboscis and its structures 

An extensible proboscis arising from the posterior of the 
rhynchocoel is present in the Drilliinae (formerly Clavinae; 
ICZN decision pending on further name change to Clavusi- 
nae) and all the radulate turrids examined, excepting Gymno- 
bela emertoni, where the radula is vestigial. A proboscis is 
present in all species of Conus, in Hastula, and in other 
radulate Terebridae, such as T. subulata, and T. babylonia 
(Taylor, 1990). The distal opening to the proboscis forms the 
true mouth as in all probosciferous gastropods. Shimek 
(1975) referred to the opening of the buccal cavity as being 
the true mouth. 

A proboscis is absent in the radula-less Turridae such as 
Teretiopsis, Taranis (Kantor & Sysoev, 1989), Philbertia 
leufroyi boothi, P. linearis (Smith, 1967, Sheridan et al., 1973) 
and the radulate Gymnobela emertoni. A proboscis is also 
absent in species of Duplicaria and Pervicacia, which are 
radulate forms of the Pervicaciidae (Taylor, 1990), and in the 
many species of Terebridae which lack a radula, such as 
Terebra maculata, T. gouldi, T. dimidiata, and T. affinis 
(Miller, 1970, 1975; Taylor, 1990). 

In Duplicaria spectabilis and Gymnobela emertoni we have 
observed a low cylinder of muscular tissue surrounding the 
opening to the buccal cavity (Fig. 8) (Taylor (1990, Fig. 2). 
We think that this may represent the remnant of a much 
reduced proboscis. A similar reduced structure found in 
Cenodagreutes spp. and Philbertia linearis, was described by 
Smith (1976) as the muscular sheath. 



130 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 

MOMBBn 




Fig. 5 Duplicaria spectabilis; relaxed, critical-point dried specimen. A, Section of the rhynchodeal wall showing the transverse muscles 
joining the inner and outer walls. Scale bar = 100 urn. B, Detail of junction of transverse muscles joining the inner wall of the 
rhynchodaeum. Scale bar = 20 um. 



Buccal tube 

The buccal tube is that portion of the alimentary canal lying 
between the buccal cavity and the true mouth, which is 
situated at the distal end of the proboscis. The buccal tube is 
present in all toxoglossans with a proboscis and is absent only 
in those species where that organ is lost. It is very short in 
Strictispira paxillus where the buccal mass lies at the extreme 
anterior end of the proboscis. 

In the Mangeliinae the epithelium of the buccal tube is very 
thin (Fig. 9), but much thicker in species of other subfami-lies 
such as the Drilliinae and Clavatulinae (Fig. 6). Shimek 
(1975) refered to the buccal lips (see below) as the buccal 
tube, and he called the true buccal tube, the inner proboscis 
wall. 



Buccal tube sphincters 

In most toxoglossans, one or more annular sphincters may be 
found in various positions within the proboscis. 



a) Distal sphincter(s) 

In most species with a proboscis, there is a distal sphincter 
around the true mouth. Frequently, there is a second sphinc- 
ter also near the proboscis tip, but located just to the 
posterior of the first (Fig. 6). In 'lower' turrids such as the 
Drilliinae Cochlespirinae and Clavatulinae, the sphincter(s) 
grip the solid, radular teeth whilst they are held at the 
proboscis tip (Sysoev & Kantor, 1989; Kantor & Taylor, 
1991). 



b) Intermediate sphincter 

A small muscular sphincter, situated about halfway along the 
length of the proboscis is found in Splendrillia (Kantor & 
Sysoev, 1989, fig. 3c). Species of Conus also have a sphincter 
situated some distance posterior to the proboscis tip (Greene 
& Kohn, 1989) which we classify as an intermediate sphinc- 
ter. 



c) Basal sphincter 

A sphincter located near the base of the proboscis has been 
described for Mangelia nebula (Sheridan et al., 1973). 
Recently, Delaunois & Sheridan (1989) have illustrated a 
section through the buccal area of M. nebula, showing a 
single radular tooth held in the buccal tube. The tooth is 
gripped at the anterior end by the buccal tube introvert (see 
below), and the posterior end by the basal sphincter (Fig. 9). 



Buccal tube introvert 

This is a muscular, flap-like structure found towards the distal 
end of the buccal tube of Mangelia nebula (Fig. 9) and called 
a valve (valvule) by Sheridan et al. (1973). Eucithara strom- 
boides has a longer, but apparently homologous structure 
(Fig. 10). Delaunois & Sheridan (1989) showed that one of 
the functions of this structure is to grip the radular tooth in 
the buccal tube, but in Eucithara where the structure is very 
long (Fig. 10), it may possibly be used to transport teeth to the 
proboscis tip. 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 

od rs 



131 




bts 



Fig. 6 Clionella sinuata;A, longitudinal section through the 
foregut; B, section of tip of proboscis showing sphincters; C, 
section of portion of the inner wall of the rhynchodeum. showing 
the differentiation in epithelium from that similar to the proboscis 
wall, to that typical of the lining of the rhynchocoel. 
Abbreviations: bm, buccal mass; bts, buccal tube sphincters; con, 
circum-oral nerve ring; mb, muscular bulb; od, odontophore; oe, 
oesophagus; p, proboscis; rhs, rhynchostomal sphincter; rs. 
radular sac; sg, salivary gland; tec, tall epithelial cells; vg, venom 
gland. 

Sac-like enlargement of buccal tube 

One other character associated with the gripping of marginal 
teeth at the proboscis tip, is the presence of a sac-like 
en-largement of the anterior or middle parts of the buccal 
tube. It is found in different 'lower' turrids (Kantor & Taylor, 
1991) as well as Mangelia nebula (Sheridan et ai, 1973) and 
Conidae (Conns catus (Greene & Kohn, 1989) and C. ventri- 
cosus). Usually, the epithelium lining the enlargement is 
formed of much taller cells than in the rest of the buccal tube. 
These cells tightly surround the single radular teeth whilst 
they are being held at the proboscis tip and may afford a 
better grip. In Splendrillia chathamensis , Sysoev & Kantor 
(1989) found the base of tooth adhering to a pad of epithelial 
cells. 

Protrusive lips of proboscis! buccal tube 

In a few species, the inner lining of the outer lips of the 
proboscis can be protruded. For example, in Turricula nelliae 
spurius, the lips (Fig. 11) are densely covered by paddle or 
discocilia, which according to Haszprunar (1985) may indi- 
cate the presence of chemosensory cells. Similar protrusible 
lips are also found in Lophiotoma leucotropis and probably in 
Aforia aulaca alaskana (Sysoev & Kantor, 1987). 

In relaxed specimens of Mangelia powisiana, a sac consist- 
ing of a single layer of cells is protruded from the proboscis 



ovg 




rhs 



sng 



Fig. 7 Conus ventricosus; longitudinal section of the foregut 
showing the proboscis retracted into the rhynchodeum. 
Abbreviations; bm, buccal mass; bts, buccal tube sphincter; dasg, 
duct of accessory salivary gland; fpw, fold of proboscis wall; ors, 
opening of radular sac; ovg, opening of venom gland; p, 
proboscis; rhs, rhynchostomal sphincter; sng, snout gland. 




5mm 



Fig. 8 Gymnobela emertoni: longitudinal section of the foregut 
showing, the remnants of the proboscis, buccal lips and vestigial 
radular sac. Abbreviations: bl, buccal lips; con, circum-oral nerve 
ring; m, mouth; pr, reduced proboscis; rhs, rhynchostomal 
sphincter; rm, radial muscles in rhynchodeal wall; rs, radular sac; 
sg, salivary gland. 



132 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



tip (Fig. 12). This sac is covered in granule-like structures 
which are formed from single cells with large rounded nuclei. 
The distinctive epithelial cells seen at the proboscis tip of 
Mangelia nebula by Sheridan et al. (1973) may be the same 
structure but in a more contracted position. The function of 
this sac structure is not known. 

Position of the buccal mass 

Three conditions are known in the Conoidea; 

a) Buccal mass situated at the base of the proboscis (Fig. 1 ) 
For three reasons we consider this condition to be the 
primitive state within the Conoidea. Firstly, a basal buccal 
mass is found in species of the subfamily Drilliinae, which 
with five teeth in each radula row, are considered to possess 
the least-derived type of radula. Secondly, and also in the 
Drilliinae, there is a muscular connection between the retrac- 
tor muscle of the radular sac and the columellar muscle 
(Kantor, 1990). This is a condition found in some meso- and 
archaeogastropods, as for example in Littorina, Cryptonatica 
and Tegula (Fretter & Graham, 1963; Kantor, unpublished 
observations). In most other probosciform gastropods, 
including those turrids where the buccal mass lies within the 
proboscis, this connection is broken and the radula is con- 
nected by muscles to the walls of the proboscis. Finally, the 
basal buccal mass is a character-state shared amongst most of 
the subfamilies of Turridae, along with the Terebridae, 
Pervicaciidae and Conidae. 

b) Buccal mass located within the proboscis 

In Clavatula diadema (Clavatulinae), the buccal mass lies 
within the proboscis, but in a proximal position (Kantor, 
1990, fig. 8). In Clionella sinuata (Clavatulinae), Pilsbryspira 
nympha (Zonulispirinae) and Funa latisinuata (Crassispiri- 
nae), it lies more anteriorly, about half way along the 
proboscis (Figs 6 & 14). In Strictispira paxillus (Strictispiri- 



nae) (Fig. 13), Toxiclionella tumida (Clavatulinae) (Kantor, 
1990 fig. 4), and Turricula nelliae spurius (Cochlespirinae) 
(Taylor, 1985; Miller, 1990), the buccal mass lies in a distal 
position near the tip of the proboscis. 

The distally shifted position of the buccal mass in these few 
turrids is a derived condition, being found only in some 
species of the subfamilies Clavatulinae, Cochlespirinae 
Zonulispirinae and Strictispirinae. 

c) Buccal mass situated a long way to the posterior of the 
proboscis base (Kantor, 1990, fig. 1). 

This condition is found only in Hormospira (Pseudome- 
latominae) and described by Kantor (1988). 

Elongation of the oesophagus to the anterior of the 
circum-oral nerve ring 

In some turrids the oesophagus is elongated into a curved 
loop between the base of the proboscis and the circum-oral 
nerve ring (Fig. 14). This elongation is found in those turrids 
with a buccal mass situated within the proboscis, and allows 
forward movement of the buccal mass on protraction of the 
proboscis. This condition is found in Clavatulinae. Stric- 
tispirinae, Turricula nelliae spurius (Cochlespirinae), Cras- 
sispirinae such as Funa latisinuata, and Pilsbryspira 
(Zonulispirinae). 

Buccal lips (inner buccal tube) 

These consist of muscular extensions of the anterior walls of 
the buccal mass, which protrude as a tube into the lumen of 
the buccal tube (Figs 1 & 9). In Oenopota levidensis where 
the buccal lips are long (Shimek, 1975), they form a second 
'proboscis' within the true proboscis. At full contraction of 
the true proboscis, the tube formed by the buccal lips 
protrudes through the mouth. Shimek (1975) called this 
secondary 'proboscis' the buccal tube. Various developments 




0-5mm 



Fig. 9 Mangelia nebula; longitudinal section through the proboscis. A, with buccal lips protracted; B, radular tooth in proboscis and buccal 
lips withdrawn into the buccal cavity. After Sheridan et al. (1973, fig. 7) & Delaunois & Sheridan (1989, plate II). Abbreviations: be, buccal 
cavity; bl, buccal lips; ds, distal sphincter of buccal tube; i, buccal tube introvert; m, mouth; ps, posterior sphincter of buccal tube; t, 
radular tooth. 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



133 




Fig. 10 Eucithara stromboides; longitudinal section through the 
anterior end of the proboscis showing the buccal tube introvert. 
Abbreviations: i, introvert; is, intermediate sphincter; m, mouth. 



The buccal cavity and radular apparatus 

From the true mouth, the buccal tube leads to a well-defined 
chamber, the buccal cavity, which is surrounded by thick 
walls of circular muscle. The radular diverticulum usually 
opens ventrally into the buccal cavity. It consists of the 
radular sac within which the radular teeth are formed, and in 
less-derived turrids, an odontophore and sublingual pouch 
(Fig. 15). The latter is the site where degeneration of the 
radular teeth and ribbon occurs. The buccal sac is defined 
(Shimek, 1976), as that part of the radular diverticulum that 
lies between the buccal cavity and the entrance of the salivary 
ducts. 

In higher turrids without a radular membrane and odonto- 
phore, the sublingual pouch is transformed into a caecum for 
the storage of radular teeth prior to their use at the proboscis 
tip. 




Fig. 11 Turricula nelliae spurius; extended proboscis, showing the 
inner ring of the protrusive lips. Scale bar = 100 um. 



of the buccal lips from a short tube to long proboscis-like 
structures, are seen in species of the subfamily Mangeliinae. 
Sections of Mangelia nebula (Sheridan et al. , 1973; Delaunois 
& Sheridan, 1989) show that the buccal lips can be inverted 
into the buccal cavity (Fig. 9b). In the genus Aforia (Cochle- 
spirinae), some species have well developed buccal lips, but 
in others they are absent (Sysoev & Kantor, 1987). 

In some conoideans lacking a proboscis and radula (e.g. 
Terebra gouldi (Miller, 1975)), the buccal lips are enlarged 
and consist only of circular muscles. They have the appear- 
ance of, and may be confused with, the true proboscis. 



Radula caecum (often called short arm of the radula 
sac) 

This is a diverticulum which branches off the anterior end of 
the radular sac, in which detached radular teeth are stored 
prior to their use at the proboscis tip (Fig. 15). We regard this 
structure as a homologue of the sublingual pouch found in 
other gastropods with a radular ribbon. A radular caecum is 
present in higher turrids, for example the subfamilies Man- 
geliidae, Daphnellinae, and Borsoniinae and also in Conidae 
and some Terebridae. 

Shimek (1976) showed that the caecum in Oenopota lev- 
idensis is divided longitudinally by a septum. We have seen 
this structure only in Micantapex parengonius (Borsoniinae). 

Radular membrane 

In general, the 'lower' turrids have a robust radular mem- 
brane, whilst in 'higher' turrids, it is thin or absent. However, 
even in 'lower' turrids, the strength of the membrane varies 
considerably between taxa and we recognise only the pres- 
ence or absence of the membrane as a functionally important 
character. 

A radular membrane is absent in the subfamilies Borsonii- 
nae, Mangeliinae, Daphnellinae, Conorbinae, Clathurelli- 
nae, Taraniinae, Conidae and most Terebridae. 

Odontophore 

An odontophore with cartilages is present in many lower 
turrids (Drilliinae, Pseudomelatominae, Strictispirinae, Clav- 
atulinae, Turrinae, Cochlespirinae, Crassispirinae), the Per- 
vicaciidae, and a few species of Hastula (Terebridae), but is 
absent in higher turrids, Conidae and most other Terebridae. 
If an odontophore is present, then the cartilages may be 
either fused, or separated at the anterior end. If the cartilages 
are separated, they are joined by a muscular connection. We 
have seen fused odontophoral cartilages in Lophiotoma, 
Pseudomelatominae, Splendrillia, Clavus sp., Inquisitor and 
Funa spp., Toxiclionella and some Aforia species. Two 
separate cartilages are usually present in species of Clavatuli- 
nae (except Toxiclionella), Strictispira paxillus (Strictispiri- 
nae) (Fig. 13). In Aforia lepta (Cochlespirinae), only the 
muscle is present, over which the radular membrane bends 
(Sysoev & Kantor, 1988). 



134 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 




Fig. 12 Mangelia powisiana; a, relaxed specimen showing sac-like structure at distal end of extended proboscis. Scale bar = 100 urn. b, 
detail of sac body with warty external surface. Scale bar = 100 ^m. c, section of the sac showing the thin epithelium with granule structures 
produced by single cells with large nuclei. Scale bar = 50 pirn, d, detail of c. Scale bar = 10 urn. 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



135 



Raclula 

The radula has been by far the most studied of the organs of 
the foregut and there are many published illustrations of 
conoidean radular teeth (e.g. Powell, 1966; McLean, 1971; 
James, 1980; Bandel, 1984; Bogdanov, 1990; Nybakken, 1990 
and Taylor, 1990). Shimek & Kohn (1981) classified turrid 
radulae into a number of functional groups and attempted a 
cladistic analysis of radular characters. However, amongst the 
'lower' turrids there is little evidence from direct observations 
to support their functional categories. Indeed, recent evi- 
dence shows that even in the least-derived radulae which 
possess a strong radular ribbon, the marginal teeth can be 
held singly at the proboscis tip in a stabbing position (Kantor 
& Taylor, 1991). 

A radula is present in most Turridae, all Conidae, possibly 
all Pervicaciidae and some Terebridae. It is absent in some 
species of Daphnellinae, Taraninae and many species of 
Terebridae (Miller, 1970; Taylor, 1990). The phenomenon of 
radula-loss in conoideans has recently been reviewed by 
Kantor & Sysoev (1989). 

For the purposes of the present analysis, we have 
attempted to recognise different morphological types of 
radula, without any functional interpretation. 

The radula of the Drilliinae, which is usually regarded as 
the least-derived condition within the Turridae, has five teeth 
in each transverse row (Fig. 16a). These teeth are usually 
refered to as central, lateral and marginal teeth respectively; 
although there are different interpretations (Kantor, 1990; 
Starobogatov, 1990). We consider the morphology of each of 
these teeth in turn. 

1. Central tooth 

A central tooth is present in species of Drilliinae, Pseudome- 

latominae, Turrinae, Clavatulinae, and Cochlespirinae. It can 



be reduced and lost in some species of these subfamilies 
except Pseudomelatominae. (i) In the Pseudomelatominae, 
the central tooth is fairly robust and broad, with a large 
curved central cusp and sometimes smaller cusps at either 
edge (Fig. 17e & f). (ii) In the Drilliinae the central tooth is 
robust, but small and narrow (Fig. 16b & d), usually with a 
prominent central cusp and a number of subsidiary cusps, (iii) 
In the Turrinae and Clavatulinae (Figs 18a-d, 19a & b), the 
central tooth appears broad, but apart from a spine-like 
central cusp is poorly defined. The central cusp appears 
homologous with the central tooth of the Drilliinae, but the 
insubstantial, lateral 'wings' may represent vestiges of lateral 
teeth which have fused with the central tooth. Alternatively, 
the whole tooth might be homologous with the central tooth 
of the Pseudomelatominae, the central cusp remaining promi- 
nent, but the lateral edges becoming less substantial. Study of 
the ontogeny of the radula in these taxa might distinguish 
between these alternative possibilities. 

2. Lateral teeth 

We recognise two types of lateral teeth, (i) In what is 
considered to be the least-derived condition, most species of 
Drilliinae have large, multicuspidate, comb-like, lateral teeth 
(Fig. 16a, c,e). However, reduced teeth are found in some 
drilliine species (Bandel, 1984, fig. 306). (ii) In Antiplanes 
(Cochlespirinae), the radula folds along the middle of the 
radular ribbon, suggesting that the poorly defined, plate-like 
teeth are in fact laterals (Kantor, 1990; Kantor & Sysoev, 
1991, figs 26-27, 30-32). These 'teeth' were not visible on 
S.E.M. preparations. Similar, poorly defined, lateral 'teeth' 
are also present in optical preparations of Crassispira and 
Crassiclava of the Crassispirinae (Maes, 1983 fig. 31 & 37, p. 
322; Kilburn, 1988, p. 239). 

In all other subfamilies of Turridae, Pervicaciidae, Tere- 
bridae and Conidae. lateral teeth are absent. 



r 



/ 




4 J. 



M£\v 



', 'rVf-l l • ;•"' 






,^ 







:-.■.&?'• *:-w\ v"r* {*££ 



Fig. 13 Strictispira paxillus; transverse section of the rhynchoel and the proboscis tip. a, mouth with distally-situated radula and virtually no 
bucal tube. Scale bar = 100 urn. b, section of the proboscis slightly to the posterior of (a) showing the two large odontophoral cartilages. 
Scale bar = 100 um. 



136 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



1mm 




Fig. 14 Funa latisinuata; anterior alimentary system. A, proboscis 
with buccal mass in extended position; B, with buccal mass in 
retracted position and showing the loop of the oesophagus 
situated to the anterior of the nerve ring. Modified from an 
unpublished drawing by J. Miller. Abbreviations: bm, buccal 
mass; bt, buccal tube; con, circum-oral nerve ring; mb, muscular 
bulb; ol, oesophageal loop; vg, venom gland. 




Fig. 15 Diagrammatic section through the radular sac. A, in 
turrids possessing an odontophore; B, turrids lacking an 
odontophore, but with a radula caecum. Abbreviations: bs, 
buccal sac; od, odontophore; rs, radular sac; re, radula caecum; 
sip, sublingual pouch; t, radular teeth. Buccal sac is that portion 
of the radular sac lying between the entrance of the salivary ducts 
and the buccal cavity. 



3. Marginal teeth 

In most conoideans the marginal teeth are the principal 
functional teeth. Although diverse in appearance, they can be 
divided into three broad categories of solid, wishbone and 
hollow. There may be several subdivisions of each category. 
Teeth of the first category are represented by a single, flat, 
distally acute plate. Wishbone teeth are characterised by two 
plates connected to each other. Hollow teeth are distin- 
guished by a cavity within the tooth. 

a) Solid marginal teeth 

We recognise four main categories of solid teeth, (i) Simple, 
flat teeth, often with a simple, blunt barb (Figs 16a, f, Fig. 
20a). This type of tooth is common in the Drilliinae. (ii) 
Simple teeth as in (i), but with the lateral edges of the tooth 
curved to form a channel or gutter. This type of tooth has 
been recorded from Drillia cydia (Powell, 1966, fig 81; Maes, 
1983, fig. 28). (iii) Simple, solid teeth, which are curved and 
pointed (Fig. 17e). This type of tooth is found only in 
Pseudomelatominae (Kantor, 1988) and the Pervicaciidae 
(Taylor, 1990). (iv) Simple, awl-shaped teeth with a large base 
and pointed tip and a spathulate process midway along the 
tooth (Figs 17a-d). This type of tooth has been found only in 
the subfamily Strictispirinae. 

b) Wishbone teeth (sometimes called duplex teeth) 

In this type of dentition, the marginal teeth consist of two 
parts, comprising the main tooth together with an accessory 
limb. Published illustrations suggest a great variety of form in 
wishbone teeth, but S.E.M. observations show that some of 
this variety results from artifacts produced by the transpar- 
ency of light microscopy and by different positions of teeth 
(often with displaced tooth parts) in preparations. 
We recognise four basic types of wishbone teeth: 
(i) Broad, slightly curved teeth, sometimes with a blunt 
barb (Fig. 20 b-d). The lateral edges of the teeth are 
thickened, with a thin accessory limb attached to the main 
tooth at the anterior and posterior ends. This type of tooth is 
common in some Crassispirinae such as Inquisitor, Paradrillia 
and Funa, where the size and shape of the accessory limb 
varies considerably between species (Kilburn, 1988). Because 
the main limb is similar to the marginal teeth of the Drillii- 
nae, we suggest this as the least-derived type of wishbone 
tooth, (ii) The teeth of this type are robust, short and curved, 
sometimes with a knife-like cutting edge on the main limb 
and a large accessory limb (Figs 18a,c; 19a, d). Teeth of this 
type are found in species of Turrinae, Clavatulinae, and 
Cochlespirinae. (iii) Teeth that may be modified wishbone 
teeth have been illustrated for Ptychobela nodulosa and 
P.suturalis by Kilburn (1989, figs 17-19). The teeth are 
awl-shaped without barbs, with apparently two nearly equi- 
size limbs joined to form a central channel. An S.E.M. study 
of these teeth is needed to claify their morphology, (iv) In the 
radula of Ptychobela griffithi the teeth appear to be robust 
and solid with a simple barb (Fig. 22a), but they may in fact 



Fig. 16 Radulae of Drilliinae. a, half radula row of Clavus sp. from Guam showing blade-like marginal teeth, comb-like lateral teeth and the 
small central tooth. Scale bar = 50 um. b, central tooth of Clavus unizonalis. Scale bar = 5 um. c, central and part of lateral teeth of 
Spirotropis monterosatoi . Scale bar = 20 um. d, central tooth of S. monterosatoi. Scale bar = lOum. e, single lateral tooth of S. 
monterosatoi. Scale bar = 20 um. f, marginal teeth of S. monterosatoi. Scale bar = 20 urn. 



Fig. 17 Radulae of Strictispirinae and Pseudomelatominae. a, radula of Strictispira paxillus. Scale bar = 50 um. b, marginal teeth of 
Strictispira stillmani. Scale bar = 50 urn. c, radula of Cleospira ochsneri. Scale bar = 50 um. d, marginal teeth of Strictispira paxillus seen 
from side. Scale bar = 50 um. e, radula of Pseudomelatoma penicillata. Scale bar = 100 um. f, central tooth of P. penicillata seen from side. 
Scale bar = 10 um. (see p. 138) 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



137 




138 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 







FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



139 




j Fig. 18 Radulae of Clavatulinae and Cochlesprinae. a, Clionella sinuata; wishbone marginal and small central teeth. Scale bar = 50 \im. b. 
Clionella sinuata small central tooth. Scale bar = 10 urn. c. Turricula nelliae spurius, radula with wishbone marginal teeth and central tooth 
with spine-like cusp and lateral flanges. Scale bar = 50 urn. d. T. nelliae spurius central tooth. Scale bar = 5 urn. 



140 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 







Fig. 19 Wishbone teeth of Turrinae and Cochlespirinae. a. radula of Gemmula deshayesi. Scale bar = 50 urn. b. marginal tooth of Gemmula 
deshayesi Scale bar = 20 jim. c. marginal tooth of Lophiotoma acuta Scale bar = 10 |xm. d. marginal tooth of Antiplanes sanctiioannis . 
Scale bar = 20 \im.. 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



141 



be formed from two pieces as in Ptychobela suturalis (see 
above). Lack of material precluded further study of this and 
the type iii wishbone teeth. 
c) Hollow teeth 

There is a great diversity of detailed variation in the form of 
hollow marginal teeth even within a single genus (see for 
example, James (1980) and Nybakken (1990) for Conus and 
Bogdanov (1990) for Oenopota). However, for the purposes 
of this analysis we recognise only five main types of hollow 
teeth, (i) Teeth of this type are long, slender, and enrolled, 



with a small base. The base is not differentiated morphologi- 
cally and is not solid. The distal end of the tooth may be 
simple, or more or less, elaborately barbed (Figs 22e,g). 
There is an opening near the distal tip and a second opening 
placed more or less terminally at the proximal end. The shaft 
oiHastula hectica is perforated by holes (Taylor, 1990, fig. 2). 
For some Conus species, Nybakken (1990) has shown that 
during ontogeny, the hollow, rolled teeth develop from open, 
guttered forms and become progressively more elaborately 
barbed. Hollow teeth of Type i are found in species of 




Fig. 20. Radulae of Clavinae and Crassispirinae. a. marginal tooth of Drillia rosacea. Scale bar = 50 (xm. b. Funa latisinuata; blade-like 
marginal teeth with thin accessory limb. Scale bar = 50 \xm. c. Vexitomina garrardi; part of blade-like marginal tooth with accessory limb 
(arrowed). Scale bar = 10 ^m. d. enlargement of (d) showing accessory limb. Scale bar = 10 \im. 



142 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



Borsoniinae, Clathurellinae, Toxiclionella (Clavatulinae), 
Conidae, and Terebridae (ii) Hollow teeth of this second type 
are often short with a large, solid base (Fig. 23). The tooth 
cavity opens laterally between the shaft and the base. There 
are frequently side projections around the base (hiked dagger 
form of Powell 1966), often with a large irregular solid 'root' 
projecting from the base (Fig. 23e,f). These teeth are often 
only partially enrolled. Barbs may be present. Marincovich 
(1973) records rows of holes in the teeth of Agathotoma 
ordinaria (Mangeliinae). Teeth of Type ii are found in the 
subfamilies Mangeliinae, Oenopotinae (Bogdanov, 1990, figs 
407-438), Thatcheriinae, and the radulate Daphnellinae. (iii) 
Teeth of this type are partially enrolled at the base, but solid 
and blade-like in the distal part (Fig. 22b). This type of tooth 
is presently known only from Hastula bacillus (Taylor & 
Miller, 1990). It may represent a transitional form between 
the solid teeth found in the Pervicaciidae and the hollow teeth 
of the Terebridae. (iv) This type of tooth is loosely enrolled 
to form a central channel, with a simple barb at the tip. The 
tooth was first described in detail from Imaclava unimaculata 
(Clavinae) by Shimek & Kohn (1981 fig. 7). Imaclava other- 
wise has comb-like lateral teeth as in typical Clavinae. Similar 
teeth are present in other species of Imaclava (McLean, 1971, 
fig. 7). (v) Enrolled teeth with a complex appearance are seen 
in Pilsbryspira nympha (Zonulispirinae) (Fig. 21). Although 
these are hollow teeth with a small barb, the shaft is complex 
and appears to be formed by partial enrolling of two units 
(Fig. 21b). The tooth may be derived by the enrolling of the 
elongate wishbone teeth typical of the Crassispirinae. (vi) 
Vestigial teeth, semi-enrolled, with a gutter along the tooth. 
Teeth of this type are considered by Bogdanov (1990) as 



derived from the distal part of the shaft of Type ii teeth. This 
type of tooth is found in Propebela turricula and P. harpularia 
(Oenopotinae) (Bogdanov, 1990, figs 41, 433). 

Glands of the foregut 

Salivary glands 

Salivary glands are present in most turrids, Conus and the 
radulate species of Terebridae and Pervicaciidae. In most 
species a pair of glands is present, but these may be fused 
together. The salivary ducts always open into either side of 
the buccal sac (Fig. 1). In Turricula nelliae spurius, which has 
a distal buccal mass, the salivary glands are contained within 
the proboscis and attached to the oesophagus (Miller, 1990). 

In most conoideans, the salivary glands are acinous, but in 
the Mangeliinae, Thatcheriinae, Daphnellinae and Hae- 
dropleura septangularis (Crassispirinae) the glands consist of 
long, convoluted, single tubes (Sheridan et a/., 1973; own 
observations). 

Turrids without a radula also lack salivary glands, but in 
the Terebridae, glands are present in some radula-less forms, 
such as Terebra gouldi and T. maculata (Miller, 1970, 1975). 

Accessory salivary glands 

These are known in a few species of Turridae, some Conidae 
(Marsh, 1971; Schultz, 1983) and Terebridae (Taylor & 
Miller, 1990; Taylor, 1990). They have a similar histology to 
the accessory salivary glands found in other neogastropod 
families such as the Muricidae (Andrews, 1991). Further- 




Fig. 21. Enrolled teeth of Pilsbryspira nympha. a. several adjacent marginal teeth. Scale bar = 25 um. b. detail of base of tooth showing 
double structure (arrow) suggesting that tooth may be formed by the enrolling of wishbone teeth. Scale bar = 5 um. 



Fig. 22 Single marginal teeth from Turridae and Terebridae. a. Ptychobela griffithi. Scale bar = 10 urn. b. Hastula bacillus. Scale bar = 
5 um. c. Glyphostoma Candida Scale bar = 50 um. d. enlargement of the tip of the G. Candida tooth. Scale bar = 10 um. e. Genota 
mitraeformis. Scale bar = 20 um. f. Terebra babylonia. Scale bar = 20 um. g. Conus ventricosus Scale bar = 20 um. 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



143 




144 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 




FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



145 



more, the ducts from the accessory glands open near the tip 
of the buccal tube, which is the homologous position to that 
found in other neogastropods. 

Within the Turridae, we have observed accessory salivary 
glands in only two subfamilies: the Borsoniinae (Scrinium 
neozelanicum, Borsonia ochracea, and Micantapex parengo- 
nius) and Cochlespirinae (Aforia hypomela, A. kupriyanovi, 
A. abyssalis). In the Terebridae, we have seen accessory 
glands in Hastula bacillus, Terebra babylonia, T. funiculata 
and T. subulata (Taylor, 1990). Usually, only a single gland is 
found, but two glands are present in Terebra subulata. 

Venom apparatus (venom gland and muscular bulb) 

The long, tubular, and convoluted venom gland is the most 
conspicuous organ of the conoidean foregut. It always passes 
through the nerve ring and always opens into the buccal 
cavity immediately posterior to the opening of the radular sac 
(Figs 1 & 7). The venom gland is present in most conoideans, 
except the radulate Strictispira (Maes, 1983); Gymnobela 
tincta, which has a vestigial radula; the radula-less turrids 
from the subfamilies Daphnellinae and Taraninae (Smith 
1967; Sheridan et al, 1973; Kantor & Sysoev, 1989), the 
radula-less Terebridae (Miller 1975; Taylor, 1990) and the 
radulate Pervicaciidae (Taylor, 1990). 

In some species, the histology of the venom gland changes 
in the anterior portion of its length, after its passage through 
the nerve ring. The posterior portion is packed with venom 
granules (Fig. 24), but the anterior portion is duct-like and 
ciliated (e.g. Clavatula, Clionella, Turricula, Lophiotoma and 
Pilsbryspira). This change in histology is usually correlated 
with the elongation of that part of the oesophagus lying 
between the nerve ring and buccal mass. In other conoideans, 
venom granules are present all the way along the length of the 
gland, sometimes even into the buccal cavity. 

Extensive studies have been made of the composition and 
pharmacology of the venom in a few Conus species (review 
by Oliviera et al., 1990). The composition of the venom is 
very complex and the results from these studies have a 
potential utility in phylogenetic analysis. However, no com- 
parable studies yet exist for the Turridae and Terebridae. 

Muscular bulb 

The muscular bulb (Figs 1 & 6) lies at the posterior end of the 
venom gland and is present in all those species possessing the 
gland. Differences between taxa are observed both in the 
number, orientation and relative thickness of the various 
muscular layers forming the wall of the bulb. The usual 
condition is of an outer, circular-muscle layer, a thin, middle 
connective tissue layer, with an inner longitudinal layer. We 
have, however, observed other configurations of the muscle 
layers. For example in Mangelia species and Eucithara, the 
outer muscular layer is very thin, but the inner layer very 
thick. Daphnella reeveana has only a single, thin muscle 
layer, whilst Conus textile has four distinct alternating circular 
and longitudinal muscle layers, three of them lying inside the 
connective tissue layer. 

Additionally, Ponder (1970) mentions that he has observed 
glandular cells in the epithelium lining the muscular bulb in 



Lucerapex (Turrinae) and Maoritomella albula (Borsonii- 
nae). We have not observed the glandular cells in any turrid 
we have examined. 

Summary of foregut anatomy 

From the foregoing discussion, it is clear that there is a great 
variety of foregut anatomy present within the Conoidea and 
considerable variation may be present even within species of 
one subfamily. As a summary, twelve of the main types of 
foregut configuration are shown diagramatically in Figs 25 & 
26. It should be emphasized that only a relatively small 
number of conoidean species have been investigated ana- 
tomically and it is likely that further types of foregut remain 
undiscovered. Nevertheless, there are several anatomical 
characters which define the Conoidea and are present in most 
representatives (and in all the least derived groups). These 
are:- 

1. The presence of a venom gland. 

2. The buccal mass located at the base of the proboscis. 

3. The proboscis formed by the elongation of the buccal 
tube. 

4. The presence of a permanent rhynchodeum. 

5. The tendency for the loss of central and lateral teeth from 
the primary five toothed radular row. 



FUNCTIONAL MORPHOLOGY OF THE 
DIGESTIVE SYSTEM AND FEEDING 
MECHANISMS IN TOXOGLOSSA 

As has been outlined in the previous section, the morphology 
of the digestive system of Conoidea and especially that of the 
Turridae, is highly varied. These variations in morphology 
probably reflect differences in feeding behaviour and diet. 
Apart from Conus, conoidean diets are still very poorly 
known. Indeed, for in excess of 4000 living species of 
Turridae, feeding information is available for less than 30 
species (reviewed by Miller, 1989). These data, derived 
mainly from gut content analysis, show that turrids feed 
mainly on errant and sedentary polychaetes and more rarely 
on other phyla such as sipunculans, nemerteans, and mol- 
luscs. Very few direct observations of the feeding process in 
the Turridae have been made (Pearce, 1966; Shimek, 1883a, 
b, c; Shimek & Kohn 1980; Miller, 1990). Because of this lack 
of information, our conclusions concerning the feeding 
mechanisms of Turridae are based upon analysis of the 
morphology of the digestive tract and by comparison with 
species whose feeding mechanism is known. 

Our classifications of feeding mechanisms is based upon 
the following characters listed in order of priority: the 
presence/absence of venom apparatus (used for immobilizing 
or killing the prey); the mode of radula function ( which may 
be used solely as a whole organ, as a whole organ with 
simultaneous use of separate teeth, or as separate teeth only 
at the proboscis tip); position of the buccal mass (either basal 
or shifted anteriorly towards the proboscis tip). We recognize 



Fig. 23 Hypodermic-type marginal teeth with a large solid bases, a. Paramontana sp. Scale bar = 2 \im. b. Propebela rugulata. Scale bar = 
10 urn. c. & d. Thatcheria mirabilis Scale bars = 20 um. e.& f. Mangelia powisiana. Scale bars = 5 \vm G. Eucithara stromboides . Scale bar 
= 10 (im. 



146 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



**" ***■. ' *H 



*%'^K - - 




*Si " 













Fig. 24 Venom gland of Clavus sp. Guam. a. section through critical-point dried venom gland showing venom granules. Scale bar = 10 \im 
b. enlargement of single venom granule. Scale bar = 1 um. 



five main and several sub-types of feeding mechanism. Some 
of these have already been described (Kantor & Sysoev, 
1990; Kantor, 1990), but are here partially revised and 
corrected. 

I. Venom gland present 

Feeding mechanism Type 1 

The first functional type of digestive system and feeding 
mechanism, that in which the radula is used only as a whole 
organ in conjunction with the venom apparatus is found 
among species of Pseudomelatominae and in Toxiclionella 
twnida (Clavatulinae) and can be subdivided into two sub- 
types. 

The first sub-type is characteristic of the Pseudomelatomi- 
nae, an endemic subfamily from western central America, 
which includes 3 genera and several species (McLean in 
Keen, 1971). The anatomy of two species Pseudomelatoma 
penicillata and Hormospira maculosa indicates the isolated 
position of the group among Conoidea (Kantor, 1988). This is 
particularly clear, from the radular morphology, which con- 
sists of a large and well developed central tooth, flanked by 
large, scythe-like, but solid, marginal teeth. 

The buccal mass is situated either at the proboscis base and 
far ahead the nerve ring in Pseudomelatoma penicillata, or in 
front of the nerve ring and distant from the proboscis base in 
Hormospira maculosa. The anterior part of the digestive tract 
forms a long curve, either by the elongation of that part of the 
oesophagus between the nerve ring and the buccal mass (P. 
penicillata), or by the elongation of the posterior part of the 
buccal tube (H. maculosa). 



Both species have a well-developed venom gland and 
although the diet of Pseudomelatominae is unknown, the 
presence of the large venom gland indicates the predatory 
mode of feeding. The gastropods also have a muscular 
proboscis with a wide oral opening but without a sphincter. 
The absence of the oral sphincter, which is usually used for 
holding single radular teeth at the proboscis tip (Kantor & 
Taylor, 1990), coupled with the curved form of the marginal 
teeth, indicate that the gastropods do not use separate teeth 
for stabbing the prey. Kantor (1988) supposed that prey 
capture occurs with the aid of the proboscis tip and is 
facilitated by the wide and highly extensible oral opening. If 
this is so, then envenomation of the prey should occur within 
the anterior part of the proboscis. This facilitates the trans- 
port of prey into the buccal cavity, by the peristaltic move- 
ments of well-developed circular muscles of the buccal tube. 

However, the presence of the elongated part of the 
oesophagus between the buccal mass and nerve ring in P. 
penicillata may indicate another mode of prey capture. In 
some turrids (e.g. Funa latisinuata. Fig. 14), the presence of 
such an elongation of the oesophagus is connected with the 
ability to evert the buccal mass, with the radula, through the 
proboscis and mouth. It is possible, that P. penicillata can 
evert the buccal mass through the mouth and use the radula 
directly in prey capture. Envenomation would in this case 
occur through the damage to the prey made by the radular 
teeth. Also the very large odontophore (the largest of all the 
turrids studied) suggests that the radula may also tear the 
prey. 

The morphology of Hormospira differs from that of 
Pseudomelatoma, in that the curve is formed by the posterior 
part of the buccal tube and elongated buccal mass. The 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 







Fig. 25 Diagram (with Fig. 26) summarizing some of the major 
types of foregut morphology found amongst the Conoidea, with 
radulae, where present, illustrated alongside. Not to scale. A. 
Clavus unizonalis; B. Clionella sinuata; C. Turricula nelliae 
spurius; D. Mangelia nebula; E. Ophiodermella biennis; F. 
Daphnella reeveana. Abbreviations: asg, accessory salivary 
glands; sg, salivary glands; rs, radular sac; vg, venom gland; black 
dots are sphincters. 

radular sac is located far behind the base of the proboscis. 
Therefore, it is doubtful that the buccal mass can be everted 
through the mouth opening. This species probably catches 
prey using the proboscis tip. Envenomation could occur 
either by the squirting of venom through the mouth, when the 
proboscis is in contact with the prey, or in the anterior part of 
the proboscis, when the prey is partly swallowed. In either 
case the radula is not used to envenomate the prey and is 
either used for further transportation in the oesophagus of for 
partial tearing of prey tissue. 

The second sub-type is found in Toxiclionella tumida and 
differs from the first in that the buccal mass is located near 
the proboscis tip (Kantor, 1990, fig. 4), which has no distal 
sphincter. This species is characterized by a radula formed of 
hollow, and barbed marginal teeth (Kilburn, 1985, fig. 14), 
which are attached all along their length to the radular 
membrane. The hollow radular teeth are similar in morphol- 
ogy to those of higher conoideans. The gastropod has a long 
venom gland and in the posterior part of the proboscis there 
is a single salivary gland with paired ducts. The radular teeth 
are sufficiently long, that during protraction of the odonto- 



iAl 



saJ 



i 



147 

H 







Fig. 26 Further types of foregut morphology found in the 
Conoidea. G. Gymnobela emertoni; H. Philbertia linearis; I. 
Conus ventricosus; J. Duplicaria spectabilis; K. Terebra subulata; 
L. Terebra maadata. 



phore, the tips would protrude through the oral opening, and 
thereby stab the prey. 

A comparable mechanism may occur in Turricula nelliae 
spurius (Taylor, 1985), which has the buccal mass located in a 
similar distal position in the proboscis to that of T. tumida, 
and during feeding can protrude the odontophore through 
the mouth opening (Miller, 1990). But T. nelliae possesses a 
sphincter in the anterior part of the buccal tube, and this 
feature usually correlates with the use of separate marginal 
teeth for stabbing (Kantor & Taylor, 1991). 

In conclusion, we suggest that a similar type of feeding 
mechanism evolved independently in Pseudomelatoma and 
Toxiclionella. In the former, the primitive character of the 
radula suggests that the feeding mechanism is primary; 
whilst in Toxiclionella it is probably a secondary feeding 
mode when compared with other members of the subfam- 
ily. It is possible that with the shift of the buccal mass to the 
proboscis tip, Toxiclionella lost the mechanism of stabbing 
the prey with single marginal teeth and instead prafrjudes 
the radula through the mouth and uses the hollow. teeth 
which remain firmly anchored to the radular membrane. ' 



148 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



Feeding mechanism Type 2 

The second feeding mechanism is typical of the majority of 
'lower' turrids and the terebrid Hastula bacillus, which pos- 
sess a well developed radular membrane and lack a radular 
caecum. The characteristic feature of this mechanism is the 
use of separate marginal teeth at the proboscis tip for 
stabbing the prey, whilst the radula is also used as a whole 
organ for different purposes (Sysoev & Kantor, 1986, 1989). 

The use of single marginal teeth at the proboscis tip by 
turrids having radulas with well developed subradular mem- 
branes has been demonstrated in representatives of all 7 
subfamilies of 'lower' Turridae (excepting the Pseudome- 
latominae) and also the terebrid Hastula bacillus (Kantor & 
Taylor, 1991). 

According to the position of the buccal mass this type may 
be divided into two sub-types. Gastropods of the first sub- 
type have the buccal mass situated at the proboscis base. 
These include species of Drilliinae, Cochlespirinae, Turrinae 
and many Crassispirinae. In these gastropods, the solid or 
wishbone marginal teeth, which become detached from the 
membrane during its degeneration in the sublingual pouch, 
are used at the proboscis tip for stabbing the prey. It should 
be noted, that separate teeth were not found in the sublingual 
pouch, therefore it does not serve for the storage of teeth. 
Meanwhile, the radula as a whole organ probably has a 
different function within the buccal cavity. This is most likely 
for the transport of food from the cavity to the oesophagus. 
Some evidence for this comes from the observations of Maes 
(1981), who noted the presence of intact sipunculans in the 
posterior part of the oesophagus of Drillia cydia (Drillinae). 
Although at first sight, it might be thought that the large, 
pectinate, lateral teeth found in this species might serve for 
tearing or rasping the prey. 

A characteristic feature of the proboscis is the presence of 
the sac-like enlargement of the anterior part of the buccal 
tube and a well-developed, distal sphincter(s). Gastropods of 
this group lack a radular caecum, so they can use only teeth 
which are sporadically detached from the membrane. Either 
the marginal teeth are not used in every feeding act, or, the 
teeth are held at the proboscis tip for a long time. That is, 
from the moment of their detachment from the subradular 
membrane to the next feeding act. We have found teeth at 
the proboscis tip in sections of 'lower' turrids much more 
frequently, than in the 'higher' turrids. Moreover, in Splen- 
drillia chathamensis , in addition to the normal buccal sphinc- 
ters of the buccal tube, teeth are attached by their base to a 
'mat' of epithelial cells in the enlargement of the buccal tube 
(Kantor, 1990, fig. 3). Such a mechanism of tooth fixation 
confirms the long-term presence of the tooth at the proboscis 
tip. Thus, the enlargement of the anterior part of the buccal 
tube, could be considered as a functional analogue of the 
radular caecum. 

The use of marginal teeth at the proboscis tip, in turrids 
with a well-developed radular membrane, explains how hol- 
low, marginal teeth might have evolved independently in 
different groups possessing the radular membrane and odon- 
tophore. For example, Imaclava (Drillinae) (Shimek & 
Kohn, 1981), has hollow teeth and most probably uses these 
at the proboscis tip for stabbing the prey in a manner similar 
to that of higher Conoidea. 

The second feeding sub-type is seen in Funa latisinuata 
(Crassispirinae), which feeds upon nemerteans. From dissec- 
tion of relaxed animals, Miller (1989, fig 6f) showed that in 



the everted position, the buccal mass with the radula is 
protruded through the mouth opening (Fig. 14a). In sections 
of animals with a contracted proboscis, the buccal mass lies 
towards the base. It is known that this species uses the 
marginal teeth at the proboscis tip (Kantor & Taylor, 1991). 
Thus, the mode of feeding may be reconstructed as follows. 
After stabbing the prey, the gastropod everts the buccal 
mass, with the walls of the buccal tube, through the mouth 
opening and picks up the prey with the protruded radula. 
With retraction of the buccal mass, the prey is pulled into the 
proboscis. Correlated with this feeding mechanism, is the 
elongation of the anterior oesophagus between the buccal 
mass and the circum-oral nerve ring. During protraction of 
the buccal mass, the oesophagus should be pulled through the 
nerve ring. But, as the nerve ring in Conoidea is highly 
concentrated, and usually tightly attached to the oesophagus, 
the only possibility is the elongation of the oesophagus itself 
anterior to the nerve ring, forming a loop, which is straight- 
ened during eversion of the buccal mass (Fig. 14b). 

In addition to Funa latisinuata, this elongation of the 
oesophagus between the buccal mass and the nerve ring has 
been found in species from several different subfamilies of 
Turridae-Pseudomelatominae, all Clavatulinae, Pilsbryspira 
nympha (Zonulispirinae), Vexitomina (Crassispirinae), Tur- 
ricula nelliae spurius (Cochlespirinae), the radulate terebrids, 
Hastula bacillus, and Pervicacia tristis (Pervicaciidae). It is 
likely, that the turrid species at least have a feeding mecha- 
nism similar to that of F. latisinuata. The elongation of the 
anterior oesophagus is usually associated with the permanent 
shifting of the buccal mass towards the distal end of the 
proboscis. This is well demonstrated in the Clavatulinae and 
probably facilitates the eversion of the buccal mass through 
the mouth. 

In all species possessing an elongated oesophagus (except 
Pseudomelatoma), there is a change in the histology of the 
ante-rior part of the venom gland after its passage through 
the nerve ring. However, such a change occurs in two species 
(Lophiotoma leucotropis and Inquisitor sp.) which lack the 
elongated oesophagus. The anterior part of the gland is 
ciliated and duct-like, with no secretory granules. This indi- 
cates, that the differentiation of the gland is connected with 
the elongation of the oesophagus and thus, the latter is a 
secondary feature. 

Feeding mechanism Type 3 

The majority of Conoidea possess the third type of feeding 
mechanism, in which separate marginal teeth are used at the 
proboscis tip for stabbing prey, and the radula not used as a 
whole organ. 

The very specialized radular morphology is the most 
remarkable and well-known feature of the toxoglossan diges- 
tive system. It is characterized by a marked tendency towards 
a reduction in the strength of the subradular membrane, 
leading to its complete absence in many species of Turridae, 
the majority of Terebridae and all Conidae. Species without a 
subradular membrane, have a radula consisting only of 
complex, hollow, marginal teeth. They are known for the 
highly specialized feeding mechanism, in which individual 
teeth are used at the proboscis tip for stabbing and killing 
prey with secretions of peptide neurotoxins produced by the 
venom gland (Oliviera et al. 1990). 

Despite the similarities with the previous feeding mecha- 
nism, those 'higher' conoideans with hollow teeth and no 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



149 



radular membrane are extremely diverse compared with the 
'lower' conoideans. Moreover, this relative diversity has 
steadily increased throughout the Cenozoic (Sysoev, 1991). 
This suggests that higher conoideans may possess some 
adaptive advantages. In our opinion these advantages lie in 
the features of the morphology of the radular diverticulum. 

The higher Conoidea lack a subradular membrane, and the 
radular diverticulum is divided into two different parts; the 
radular sac and radular caecum (also known as long and short 
arms). The caecum serves for the storage of the fully-formed, 
marginal teeth. Many teeth can be stored; for example, in a 
specimen of Mitromorpha (Mitrolumna) sp. there were 106 
teeth in the radular sac compared with 64 in the caecum 
(Kantor & Sysoev, 1990). Species of higher Conoidea can 
probably use several teeth in each feeding act. For example, 
observations on the feeding of Conus textile showed that up to 
17 teeth can be used in the same attack (Schoenberg, 1981). 
By contrast, in lower turrids, there is no caecum and probably 
no more than a single tooth can be used in each feeding act. 
Predatory attacks by higher Conoidea are thus likely to be 
more successful, and the mechanism of prey capture probably 
more efficient. This may explain the relative success of the 
higher Conoidea. 

The feeding and diets of gastropods of this functional type 
are well known (Oliviera et al. 1990) and it is unnecessary to 
describe the process in detail. Only the most important 
morphological features should be noted. These are the vesti- 
gial, or completely reduced, radular membrane; the absence 
of an odontophore; the presence of a radular caecum where 
the fully-formed marginal teeth are stored, and a well- 
developed oral sphincter for gripping the teeth. The radula is 
represented by hollow, marginal teeth. The tooth ligament 
(long flexible stalk attached to the tooth base) is probably the 
rudiment of the radular membrane (Fig. 23c). Also the 
gastropods of this group often have enlarged rhynchostomal 
lips. In some species, the lips are able to invert (i.e. to form 
an introvert or pseudoproboscis) and this is also used in prey 
capture. It should be noted, that some vermivorous species of 
Conus (Marsh, 1970) and the fish-feeding C. geographus 
(Johnson & Stablum, 1971) do not stab they prey in every 
feeding act. This is possibly an initial stage of transition to 
feeding mechanism type 5. 

Usually, teeth are gripped at the proboscis tip by the buccal 
tube sphincter, but in some turrids the buccal tube introvert 
(valvule of Sheridan et al., 1973) is involved (Fig. 9). This 
structure has been reported so far in Mangelia nebula 
(Sheridan et al., 1973; Delaunois & Sheridan, 1989) and in 
Eucithara stromboides (Fig. 10). It is also possible, that the 
buccal tube introvert can be everted through the mouth 
opening and have a role in holding the prey. 

After envenomation, the prey may be held by the tooth 
itself, as occurs in many vermivorous species of Conus 
(Kohn, 1959), or with the mouth. The buccal lips may play a 
role in the transport of prey to the buccal cavity. These are 
highly protrusive in many Mangeliinae, and at least in M. 
nebula (Fig. 9) can be retracted into the buccal cavity 
(Delaunois & Sheridan, 1989). A similar possibility was 
described for Oenopota by Bogdanov (1990), who suggested 
that the buccal lips and the proboscis itself might be inverted 
into the buccal cavity. 



II. Venom gland absent 

Feeding mechanism Type 4 

Gastropods of this group have a radula with a well-developed 
radular membrane and a proboscis may be either present or 
reduced. According to the position of the buccal mass they 
can be divided into two sub-types. 

Conoideans of the first sub-type which at present includes 
only Strictispira and probably Cleospira, have the buccal mass 
located at the tip of a well-developed proboscis (Fig. 13). The 
buccal mass and radular apparatus are large, with two large 
odontophoral cartilages and massive odontophoral and pro- 
boscis retractor muscles. The radula has a strong membrane 
with two rows of solid, awl-shaped, marginal teeth. The 
buccal tube is very short and there are no oral sphincters. 
Apart from the record of polychaete setae in two individuals 
of Strictispira paxillus (Maes, 1983), nothing is known of the 
habits of this group. 

The terminal position of the buccal mass on the muscular 
proboscis, the short buccal tube and the massive radular 
apparatus, suggest that when the gastropod is feeding the 
radula is protracted out of the extended proboscis tip. The 
solid teeth and absence of venom apparatus suggest that the 
radula is involved in biting and tearing rather than stabbing. 
The feeding mechanism is thus probably more similar to 
other neogastropods such as the Buccinidae rather than to 
other conoideans. 

Conoideans of the second sub-type differ from these of the 
first one in possessing a basal buccal mass. The radula is 
well-developed, whilst the proboscis is either absent or highly 
reduced, and a rhynchostomal introvert is usually present. 
This feeding mode is found in the Pervicaciidae. The diet of 
this family is largely unknown, except for 'Terebra' nassoides 
which feeds on capitellid polychaetes (Taylor, 1990). 

In the Pervicaciidae, the absence of a proboscis means that 
the rhynchodeal introvert becomes the main organ of prey 
capture, as occurs in some proboscis-less terebrids such as T. 
gouldi (Miller, 1975). Prey are presumably pulled into the 
rhynchocoel by the introvert. In Duplicaria spectabilis there 
are large muscular buccal lips and probably a protrusive 
odontophore (Taylor, 1990, fig. 7). However, in Pervicacia 
tristis and Duplicaria kieneri there is a septum with a narrow 
aperture dividing the rhynchocoel and it is very unlikely that 
the odontophore can be protruded through the septum. 
Although we have no direct evidence, it is possible that the 
septum functions to hold prey during swallowing and perhaps 
early digestion. 

Feeding mechanism Type 5 

Finally, there are many conoideans which lack a radula, 
venom and salivary glands. Gastropods of this group include 
some Daphnellinae, Taraninae and some Terebridae. In 
addition to the absence of foregut glands and radula, a 
characteristic feature of these species is the very reduced size 
or complete absence of the proboscis. Radula-less Conoidea 
either have well-developed, rhynchostomal lips or a large 
rhynchostomal introvert, as for example, in the Terebridae 
(Miller, 1975) or Philbertia linearis (Sheridan et al., 1973). It 
is possible, that a rhynchostomal introvert is also present in 
Teretiopsis, although all sectioned specimens have it in the 
extended position and it was overlooked during the original 
description (Kantor & Sysoev, 1989). In some turrids, such as 



150 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



Cenodagreutes (Smith, 1967) and Abyssobela atoxica (Kantor 
& Sysoev, 1986), which lack the rhynchodeal introvert, there 
is a vast rhynchocoel and well-developed cavity between the 
rhynchodaeum and body walls. The walls of this cavity are 
connected by numerous transverse muscles. Both the intro- 
vert and cavity are lacking in the genus Taranis (Taraninae). 

A feeding mechanism for radula-less species is known for 
some terebrids (Miller, 1970, 1975). Thus, Terebra gouldi 
which has a relatively short introvert feeds upon the enterop- 
neust Ptychodera flava, and Terebra maculata with a long 
introvert feeds on polychaetes. Prey are caught with the aid 
of the introvert. Turrids lacking the introvert, but with the 
cavity between the rhynchodaeum and the body walls, prob- 
ably engulf prey by contraction of the radial muscles in the 
wall. This would cause negative pressure and an increase in 
the inner volume of the rhynchocoel. 

The origin of the radula-less feeding mechanism can be 
easily envisaged. It is known, that in some Conus species 
hypodermic envenomation is not necessary in each feeding 
attack (Kohn, 1959; Marsh, 1970; Sanders & Wolfson, 1961). 
It is probable that some Turridae and Terebridae, especially 
those with well-developed rhynchostomal lips or introvert, 
also feed without stabbing the prey with radular teeth. Thus, 
Daphnella reeveana, which possesses a venom gland, has a 
very short proboscis and is probably unable to hold a tooth at 
its tip (Fig. 4). As stabbing of the prey becomes unnecessary, 
the proboscis, venom gland and radula disappear. An inter- 
mediate stage is found in Gymnobela emertoni, in which the 
proboscis and venom gland have disappeared, but there is 
still a very short and reduced radular sac, opening to the 
outer side of the buccal lip (Fig. 8). 



RELATIONSHIPS OF THE CONOIDEA 



Monophyly of the Conoidea 

There has been much discussion concerning the relationships 
of the Conoidea to other prosobranch gastropods; some 
considering them to be part of a monophyletic group with 
other neogastropods (Ponder, 1973; Taylor & Morris, 1988), 
whilst others suggest an origin entirely independent of the 
neogastropods (Sheridan et al. 1973; Shimek & Kohn, 1981; 
Kantor, 1990). 

In this section we briefly review some of the evidence for 
the relationships of the Conoidea with other prosobranchs. 
Some of this evidence has been discussed in some detail by 
Kantor (1990) and only the principal arguments are presented 
here. 

The location of the buccal mass at the base of the proboscis 
as found in most conoideans, is different from the situation 
seen in most neogastropods, where the buccal mass is found 
at the distal end of the proboscis. The proboscis in most 
conoideans is formed by the elongation of the buccal tube, 
whilst in neogastropods it originates from the elongation of 
the anterior oesophagus (Ponder, 1973). However, a basal 
buccal mass is now known for the neogastropod Benthobia 
(Pseudolividae) which also exhibits a number of other primi- 
tive characters, and in Amalda (Olividae) (Kantor, 1991). 
Additionally, in Benthobia, the radular retractor muscle 
passes through the nerve ring and is connected to the 
columellar muscle (Kantor, 1991 fig. 15a). This condition is 
seen species of the turrid subfamily Drilliinae, and in most 



lower caenogastropods, but is absent in probosciform caeno- 
gastropods. 

A key autapomorphy of the Conoidea is the possession of 
the venom apparatus, comprising the venom gland and 
muscular bulb. There has been much discussion concerning 
the homology of this gland. But, Ponder (1970; 1973) 
showed, that in the neogastropod family Marginellidae a long 
coiled gland, similar in general appearance to the conoidean 
venom gland is formed by the stripping off of glandular folds 
from the oesophagus. In some marginellids the gland termi- 
nates at the posterior in a muscular bulb which is homologous 
with the gland of Leiblein. The venom gland of conoideans 
may have been derived in a similar way and is probably 
homologous with the glandular folds of the oesophagus and 
the gland of Leiblein in other neogastropods. 

The possession of tubular, accessory salivary glands is also 
considered to be an apomorphy of the Neogastropoda (Pon- 
der, 1973). These glands are patchily distributed amongst 
conoideans, but are known in some Turridae, Conidae and 
Terebridae. Both the histology of the glands (Schultz, 1983; 
Andrews, 1991) and the position of the opening of the ducts, 
confirms their homology in the Conoidea and in other 
neogastropods. The primitive Benthobia also has a large 
accessory salivary gland (Kantor, 1991). 

A radula with five teeth in each row, as is found in the 
turrid subfamily Drilliinae, has been considered as evidence 
for a separate origin of the Conoidea and Neogastropoda, the 
latter normally have three or less teeth in each row. (Shimek 
& Kohn, 1981). However, it is now known that some Olivella 
and Nassariidae have five teeth in each row (Bandel, 1984; 
Kantor, 1991). All this suggests is that the common ancestor 
of the Conoidea and the other neogastropods possessed five 
or more teeth in each row. 

In conclusion, conoideans share a number of characters 
with the neogastropods which suggest a common ancestry. 
Nevertheless, the evidence both from the position of the 
buccal mass and the formation of the proboscis, suggests an 
early divergence of the two groups. An evolutionary scheme 
for the derivation of the conoidean intraembolic proboscis 
from the acrembolic type, typical of many mesogastropods, 
has been developed by Kantor (1990). His arguments cor- 
roborate and elaborate Ponder's (1973) hypothesis that the 
Conoidea diverged from the other neogastropods before the 
formation of the proboscis. Ontogenetic studies of proboscis 
and foregut development in the Conoidea and other neogas- 
tropods might provide corroborative evidence. 

Relationships within the Conoidea 

Phylogenetic analysis 

We attempted to determine relationships within the 
Conoidea using cladistic analysis of many of the foregut 
characters described in the first part of this paper, combined 
with a few shell characters. 

Taxa used 

We have included 40 species in the analysis, with at least one 
from all the currently-recognised, subfamilies. In a few cases 
we have used previously published work. The species studied 
represent only a small proportion of living species from any of 
the subfamilies. Some of these subfamilies are very diverse 
and morphologically disparate and our sample is certainly 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



151 



Table 2. Characters and character states of the foregut and shell used in cladistic analysis. See text for details of foregut characters. 
* denotes characters where the states were treated as unordered. 



Foregut characters 

1. Rhynchodeal introvert 
*2. Rhynchodeal sphincter 

3. Accessory proboscis structure 

4. Proboscis 

5. Transverse muscles in rhynchodeum wall 

6. Epithelium of posterior rhynchodeal 
wall continuous with proboscis wall 

*7. Sphincter at distal end of buccal tube 
8. 

Sphincter in middle of buccal tube 
9. Sphincter at base of buccal tube 

10. Buccal tube introvert ('valvule') 

1 1 . Protrusive lips of buccal tube 

12. Position of buccal mass 

13. Connection of radular retractors 
to columellar muscle 

14. Extensible buccal lips 

15. Septum dividing anterior and 
posterior areas of the rhynchocoel 

16. Elongation of oesophagous between 
buccal mass and nerve ring 

17. Salivary glands 

18. Salivary ducts 

19. Type of salivary gland 

20. Accessory salivary glands 

21. Radula 

22. Radular caecum 
*23. Central tooth 

24. Lateral teeth 

25. Marginal teeth 

*26. Type of solid radular teeth 
*27. Type of wishbone teeth 
*28. Type of hollow teeth 
29. Venom gland 

*30. Connective tissue layer of muscular bulb 

*31. Muscle layers of muscular bulb 

32. Odontophore 

33. Odontophoral cartilages 

Shell and opercular characters 

*34. Shell form 

*35. Number of protoconch whorls 

*36. Sculpture of the protoconch 

*37. Siphonal canal 

*38. Position of the anal sinus 

39. Presence of apertural ornament 

(teeth on the outer lip) 

*40. Number of the teleoconch whorls 

*4 1 . Development of subsutural ramp 

42. Operculum 

43. Position of opercular nucleus 



— absent, 1 — present 

— present anterior, 1 — present posterior, 2 — absent 

— absent, 1 — present 

— present, 1 — absent 

— absent, 1 — present 

— absent, 1 — present 

— absent, 1 — one sphincter, 2 — two sphincters 

— absent, 1 — present 
— absent, 1 — present 
— absent, 1 — present 
— absent, 1 — present 
— basal, 1 — distally shifted 

— present. 1 — absent 
— absent, 1 — present 

— absent, 1 — present 

— absent, 1 — present 

— two/one glands present. 1 — glands absent 

— two ducts present, 1 — one duct present 

— acinous, 1 — tubular 

— two/one glands present. 1 — glands absent 

— present, 1 — absent 

— absent, 1 — present 

— robust muriciform, 1 — narrow 2 — broad with central spine 

— comb-like, 1 — absent 

— solid, 1 — wishbone, 2 — hollow, 3 — absent 

— Hat, 1 — curved-pointed, 2 — semi-enrolled (Hastula bacillus) 

— large blade, small accessory limb, 1 — short knife type, equilimbed 

— large base, 1 — thin small base 

— present, 1 — present with changed histology in anterior portion, 2 

— absent 

— present, 1 — absent 

— more or less equal, 1 — outer layer thin, 2 — single layer only 

— present, 1 — absent 

— not fused, 1 — fused 



— fusiform, 1 — coniform, 2 — turreted. 3 — terebriform, 4 

rounded 

— less than two, 1 — more than two 

— absent or very weak, 1 — present 

— not differentiated. 1 — moderate, 2 — long 

— sutural, 1 — shoulder, 2 — peripheral, 3 weak or absent 

— absent, 1 — present 

— less than 4, 1 — from 4 to 8, 2 — more than 9 

— absent, 1 — present 

— present, 1 — absent 

— terminal, 1 — mediolateral 



inadequate. Although anatomical data are available for many 
terebrids (Taylor, 1990 and unpublished), most of these were 
eventually excluded from the analysis for the following rea- 
son. Many of the morphological trends in the Terebrinae, 
involve partial to total loss of the foregut organs (Taylor, 
1990); thus many of the characters used in the cladistic 
analysis were recorded as missing. In our earlier attempts at 
cladistic analysis, terebrid species tended to appear in rather 
disparate positions on the cladograms. Consequently, we 
have used only three species to represent the Terebrinae and 



Pervicaciinae, the taxa being the least-derived known for 
each group. 

Characters 

We used 43 characters, coded as 101 states in the analysis. Of 
these, 35 characters concerned foregut anatomy and a further 
eight, the shell or operculum. The characters and their states 
are listed in Table 2. Full discussion of the anatomical 
characters will be found in the section of this paper concern- 



152 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



ing foregut anatomy. Additionally, brief descriptions of the 
shell characters used are given in Appendix 1. 

Outgroup 

The relationships of the Conoidea to other Neogastropoda 
are very unclear and there is no obvious sister group. In our 
various analyses we used two outgroups. The first is Bentho- 
bia the most primitive non-coniodean neogastropod known 
(Kantor, 1991). This gastropod has a buccal mass situated at 
the base of the proboscis, a muscular connection between the 
radular retractors and columellar muscles, and a full set of 
glands connected with the oesophagous. Additionally, we 
used as a second outgroup a hypothetical ancestral taxon 
consisting of the underived states, where known, of all the 
characters used in the analysis. 

Method 

The data were analysed using version 3.0 of the PAUP 
program (Swofford, 1991). Characters 2, 7, 23, 25, 26, 27, 28, 
29, 30, 31, 33, 34, 35, 36, 37, 38, 41, 42 were treated as 
unordered. The matrix of taxa and character states is shown 
in Table 3. 



Results of phylogenetic analysis 

Although we have used many new anatomical characters, the 
cladistic analysis gave rather disappointing results. The taxa 
of 'lower conoideans' especially, were rather poorly resolved 
with major branches supported by rather few weak charac- 
ters. Additionally, small adjustments to the data set produced 
rather large changes in tree topography and the number of 
alternative trees generated. 

Despite these limitations we thought it worthwhile to 
present the results of our analysis, which is the first for the 
Conoidea to use anatomical characters. Future work will 
extend on the character set shown in Table 3 and hopefully 
improve the resolution of the analysis. 

A heuristic search with the matrix shown in Table 3 and 
with Ancestor as outgroup, produced over 900 equally parsi- 
monious trees (189 steps; consistency index = 0.296; 
homoplasy index 0.704). A 50% majority rule consensus 
cladogram derived from these trees is shown in Fig. 27. 
Despite the large number of trees generated, most of the 
trees are very similar to each other and most of the branches 
are supported in 75-100% of the trees. Autapomorphies of 
the internal nodes are listed in Table 4. 

The least-derived group are the two species of Drilliidae, 



Table 3. Matrix of taxa and character states used in the analysis. See Table 2 for further details of characters and caption to Fig. 27 for taxon 
abbreviations. 



Ancestor 











(1 


7 





(1 























II 


(1 


1) 


II 


II 


(1 


II 


(1 


9 


n 


(I 


7 


7 


7 


(l 











1) 


7 


1 


7 


1 


3 





7 


7 





Benthobi 





1 











1 





























1 











1 








7 


1 





1 


7 


7 


2 


7 


7 





1 


4 











3 














PseudomP 

















1 




















1 








1 











1 











1 





1 


7 


7 

















4 


1 





1 


1 





1 


1 





StrictiP 























7 


7 








1 


1 








1 











1 








7 


1 





1 


7 


7 


2 


7 


7 








4 











1 





1 


1 





ClavusUn 




















2 






































1 








1 











7 


7 














1 


4 











1 





1 


1 





SplendrC 




















1 


1 



































1 








1 











7 


7 

















4 











1 





1 








ClavatuD 





2 











1 


1 

















1 








1 











1 








2 


1 


1 


7 


1 


7 














1 











1 


1 





1 


1 


1 


ClavatuC 

















1 


1 














1 


1 








1 


1 








1 








2 


1 


1 


7 


1 


7 


1 











1 











1 


1 





2 


1 


1 


ClionelS 

















1 


2 














1 


1 








1 











1 








2 


1 


1 


7 


1 


7 


1 











1 


4 











1 





1 


1 


1 


ToxicliT 

















1 

















1 


1 


1 





1 











1 








7 


1 


2 


7 


7 


1 

















4 








1 


1 





1 


1 


1 


LophiotL 





1 














1 











1 





1 




















1 








7 


1 


1 


7 


1 


7 


1 























2 


2 





2 








PolystiA 





1 














1 

















? 




















1 








7 


1 


1 


7 


1 


7 














7 








1 


2 


2 





2 








TurricuN 

















1 


2 











1 


1 


1 


1 





1 











1 








2 


1 


1 


7 


1 


7 


1 























2 


1 





2 


1 


1 


AforiaAb 

















7 


1 

















1 


1 


























2 


1 


1 


7 


1 


7 





1 














7 


7 


2 


1 





1 








FunaLati 





1 











1 


2 

















1 


1 





1 











1 








7 


1 


1 


7 





7 


1 











1 


4 








1 


1 





2 


1 





VexitomG 

















1 


1 


1 














1 








1 











1 








9 


1 


1 


7 





7 


1 











1 


4 


1 





1 


1 





1 








PilsbryN 

















1 


1 














1 


1 


1 





1 











1 








7 


1 


2 


7 


7 


1 


1 











1 


4 


1 








1 





1 


1 





MicantaP 




















1 

















1 




























7 


1 


2 


7 


? 
















7 


4 








1 


2 





1 








BorsoniO 





1 








1 





1 

















1 

















1 










? 


1 


2 


7 


7 


1 













7 











1 


1 





1 


1 





TomopleV 





2 














2 

















1 











1 


7 


7 


1 







7 


1 


2 


7 


7 


1 













7 


4 


1 


1 





1 





2 








TropidoF 





1 














1 

















1 


1 











1 





1 







7 


1 


2 


7 


7 
















7 


2 








1 


1 





1 


1 





Ophiodel 





1 














1 

















1 


1 

























7 


1 


2 


7 


9 


1 








1 




? 











1 


1 





1 





1 ? 


AnarithM 





1 











1 


? 


7 














1 














7 





1 







7 


1 


2 


7 


7 
















7 


4 


1 








3 


1 








1 ? 


GlyphosC 





1 














7 


7 














1 














1 





1 







7 


1 


2 


7 


7 


1 













7 


4 


1 





1 


1 


1 


1 





1 ? 


EucithaS 























1 





1 


1 





1 

















1 


1 







7 


1 


2 


7 


7 
















7 


4 


1 


1 





1 


1 


1 


1 


1 ? 


MangeliN 

















1 


1 





1 


1 


1 





1 


1 














1 


1 







? 


1 


2 


? 


7 











1 




7 


4 


1 


1 


1 


1 





1 





1 ? 


MangeliP 

















1 


1 








1 


1 





1 


1 














1 


1 







7 


1 


2 


7 


7 











1 




7 


4 


1 


1 


1 


1 





1 





1 ? 


OenopotL 





1 














7 

















1 


1 

















1 







? 


1 


2 


7 


7 











1 




7 


4 





1 


1 


1 





1 








PhilberP 













1 

















1 





1 


1 


1 











1 


1 







7 


1 


2 


? 


7 








7 


7 




7 


4 


1 


1 


1 


3 


1 


1 





1 ? 


PhilberL 
















? 


7 


7 


7 











1 


1 








1 


7 


? 


1 


1 




7 


1 


7 


7 


7 


? 


2 


7 


? 




7 


4 


1 


1 


1 


3 


1 


1 





1 ? 


DaphnelR 










1 





1 




















1 


1 


1 








7 


1 


1 


1 




7 


1 


2 


7 


? 








1 


2 




7 


4 


1 


1 








1 


1 





1 ? 


GymnobeE 










1 


1 





7 


7 


7 











1 


1 














1 


1 







7 


1 


2 


7 


7 





2 


7 


? 




7 


4 


1 


1 


1 








1 


1 


1 ? 


TeretioL 










1 


1 





7 


? 


7 











1 











1 


7 


7 


1 


1 




7 


1 


7 


7 


7 


? 


2 


7 


? 




7 


4 


1 


1 


1 











1 


1 ? 


AbyssobA 











1 


1 





? 


7 


7 











1 


1 








1 


? 


? 


1 


1 




7 


1 


7 


7 


? 


7 


2 


7 


? 




? 





? 


7 


2 











1 


1 ? 


Benthofa 





1 

















1 














1 




























7 


1 


2 


7 


7 


1 













7 





7 





1 








1 








GenotaNi 























1 














1 


7 

















1 







7 


1 


2 


? 


7 


1 













? 





1 





1 


1 





1 


1 


1 ? 


ThatcheM 





1 
































1 

















1 


1 







7 


1 


2 


? 


7 








1 


2 




7 


2 


1 


1 


1 








1 





1 ? 


TaranisM 





2 





1 








? 


7 


? 











1 


1 








1 


? 


? 


1 


1 




7 


1 


7 


7 


7 


? 


2 


? 


7 




? 


4 





1 


1 


1 











1 ? 


ConusVen 














1 








1 














1 


1 











1 













7 


1 


2 


7 


? 


1 













7 


1 


1 








3 





1 








PervicaT 


1 








1 








7 


? 


7 











1 


1 


1 


1 











1 





7 


? 


1 





1 


? 


? 


2 


? 


? 





1 


3 











3 





1 








HastulaB 


1 





1 





1 





1 

















1 





























7 


1 





7 


7 


7 

















3 


1 








3 





2 








DuplicaC 


1 


2 


1 


1 








7 


7 


? 











1 


1 


1 














1 








7 


1 





1 


7 


? 


1 


7 


? 








3 











3 





2 









FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



153 



Table 4. Synapomorphies for interior nodes. Nodes numbered as 
in Fig. 27. 



Node 



Synapomorphies (Character: state) 



1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 



7(1), 20(1), 35(0), 37(0), 38(1) 

13(1), 23(2), 24(1), 26(1) 

1(1), 3(1), 34(3), 38(3), 40(2) 

4(1), 14(1), 15(1), 29(1) 

25(1), 37(1) 

6(1), 16(1), 41(1) 

7(0), 23(0), 25(0) 

12(1) 

33(1), 43(1) 

7(2), 29(1) 

12(1) 

34(0), 40(0) 

14(1), 27(0), 43(0) 

7(1), 35(1) 

34(0), 37(2) 
2(1), 38(2), 40(2) 
22(1), 25(2), 32(1) 

20(0) 

2(1), 34(0) 

2(1), 14(1), 28(0) 

18(1) 

14(0), 35(1), 39(1), 42(1) 

7(1), 35(1) 

8(1), 34(0) 

20(0), 39(0) 

19(1), 36(1) 

28(0), 42(1) 

10(1), 11(1) 

6(1), 7(1), 14(1), 31(1) 

30(1), 31(2), 38(0) 

1(1), 14(1), 21(1), 39(1) 

38(3) 

4(1) 

5(1), 29(2), 39(0), 41(1) 

17(1), 40(0) 

1(0), 35(0) 



which are the only conoideans possessing five teeth in each 
radular row. They also retain the connection of the radular 
retractor muscle to the columellar muscle. Their distinctive 
apomorphy is the possession of large, comb-like lateral teeth. 
We have studied only three species in this group (the third 
species identical to Clavus unizonalis) which are very similar 
to each other. However, we note the very different hollow, 
enrolled 'hypodermic-style' marginal teeth of lmaclava 
(Shimek & Kohn, 1981) and the possible 'wishbone' margin- 
als of Drillia roseola (McLean, 1971). Anatomical studies of 
these taxa are needed to determine their status. 

All other conoideans are separated from the Drilliidae at 
Node 2 by the loss of the radular retractor/columellar muscle 
connection, by the loss of the lateral teeth and possession of 
curved pointed marginal teeth. None of the non-drilliid taxa 
that we have included in the cladistic analysis possess lateral 
teeth, although what appear to be vestigial lateral teeth are 
seen for example in Antiplanes (Kantor & Sysoev, 1991) and 
a few other species. Also, it is possible that the broad central 
teeth seen in Cochlespirinae may be formed by fusion of 
lateral teeth. Another apomorphy at this node is the posses- 
sion of a broad central tooth with a spine-like central cusp. 



Node 3 separates the Terebridae, with five apomorphies 
including the possession of a rhynchodeal introvert and the 
accessory proboscis structure. The Pervicaciinae (Node 4) are 
separated from Hastula (representing the Terebrinae) by the 
loss of the proboscis, the presence of extensible buccal lips, a 
septum in the rhynchocoel (although this is present in some 
Terebrinae) and the loss of the venom gland. 

Node 5 separates all other conoideans with two apomor- 
phies namely the presence of wishbone marginal teeth and a 
moderately long siphonal canal. The latter is a weak charac- 
ter and although we consider the fomer to be a strong 
character, some taxa in Clade 6 have solid teeth which PAUP 
considers a reversal from the wishbone condition. 

Clade 6 comprises taxa with the epithelium of the posterior 
part of the rhynchodeum continuous with that of the probos- 
cis and with an elongated loop of oesophagus anterior to the 
nerve ring. 

Clade 7 includes two taxa with solid marginal teeth and no 
buccal tube sphincter and Toxiclionella which has hollow 
teeth. PAUP treats the solid teeth as a reversal, but we think 
that this is unlikely. However, it is possible that the 'flanges' 
on the teeth of Strictispirinae may be modifications of a 
second limb on the tooth. Toxiclionella and Strictispira are 
grouped together at Node 8, because both have a buccal mass 
situated at the distal end of the proboscis. However, Toxi- 
clionella shares many characters with the Clavatulinae 
(including the medio-lateral nucleus of the operculum), but 
has a very different radula with hollow and barbed marginal 
teeth firmly attached to the radular ribbon located in the 
distal buccal mass. Although Toxiclionella tumida lacks a 
central tooth, a clavatuline type central is known in T. elstoni 
(Kilburn, 1985). Turricula nelliae (Node 12) shares many 
apomorphies with clavatuline species and should be trans- 
fered from the Cochlespirinae to the Clavatulinae. 

PAUP suggests that Funa and Vexitomina (Crassispirinae) 
and Pilsbryspira (Zonulispirinae) are derived from the Clav- 
atulinae. They share a number of characters, but Funa and 
Vexitomina have distinctive wishbone teeth with one broad 
flat limb and a small, thin, subsidiary limb. Pilsbryspira has 
enrolled marginal teeth and a distal buccal mass. This type of 
tooth could be derived by enrollment of the crassispirine type 
of wishbone tooth. Both groups have an operculum with a 
terminal nucleus which PAUP treats as a reversal from the 
medio-lateral nucleus of the Clavatulinae. 

Lophiotoma and Polystira (Turrinae) (Node 16) have a 
peripheral anal sinus and a posteriorly situated rhynchodeal 
sphincter. Aforia has an accessory salivary gland and PAUP 
treats this appearance as a reversal, the glands having already 
been lost between the outgroup and the first node. However, 
it is highly unlikely that these glands are regained once lost. 
Accessory glands have a very patchy distribution amongst the 
Conoidea (Conus, Benthofascis and some Clathurellinae) and 
apart from their occurrence in some terebrids, Aforia is the 
only 'lower' conoidean in which we have seen the glands. The 
distribution of this character should become clearer as more 
species are examined. Maybe significant, is the fact that 
Aforia is the only other conoidean in which the multidigitate 
osphradial leaflet typical of Conus has been found (Sysoev & 
Kantor, 1988 fig. 2J). 

From Node 18 onwards are all the so-called 'higher' 
conoideans, which in all our analyses form a monophyletic 
group. The apomorphies which define this node are the 
presence of a radula caecum for storage of detached radular 
teeth, hollow, enrolled marginal teeth, loss of the radular 



154 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 




Ancestor 
C lav us U 
Splendnllia C 



Hastula B 
Pervicacia T 
Ouphcana C 
Pseudomelatoma 
Toxichonella T • 
Strict is pir a P — 
Clavatula D 
Clionella C 
Clavulata S 
Turncula N 
Funa L 

Vexitomina G 
Pilsbryspira N 
Aforia A 



Drilliidae 
Terebridae 



Pseudomelatomidae 

Clavatulinae 

Stnctispindae 

Clavatulinae 



Lophiotoma L 
Polystira A 
Micantapex P 
Borsonia 
Ophiodermella _ 
Oenopota L — 
Tropidoturns F 
Ananthma M 
Glyphostoma C — 
Genota N — i 
Benthofascis _J 
Conus V 

Tomopleura V — 
Eucithara S 
Mangelia N 
Mangelia P — 
Thatcheria M 
Philbertia P 
Philbertia L 
Daphnella R 
Gymnobela E 
Teretiopsis L 
Abyssobela A 
Taranis M — 



Crassispirmae 

Zonulispirmae 
— Cochlespinnae 

Turnnae 

Clathurelhnae 

• Oenopotinae 
Clathurelhnae 

Conorbinae 

Coninae 

Clathurelhnae 

Mangehinae 



Daphnelhnae 



Taraninae 

Fig. 27 Majority-rule (50%) consensus tree. Autapomorphies for each node given in Table 4. Higher taxa names at the top of branches 
reflect our new classification. Taxon abbreviations in order top to bottom on the tree: Clavus U = Clavus unizonalis, Splendrillia C = 
Spendrillia chathamensis , Hastula B = Hastula bacillus, Duplicaria C = Duplicaria colorata, Pseudomelatoma P = Pseudomelatoma 
penicillatus, Toxichonella T = Toxichonella tumida, Strictispira P = Strictispira paxillus , Clavatula D = Clavatula diadema, Clionella S = 
Clionella sinuata, Clavatula C = Clavatula caerulea, Turricula N = Turricula nelliae, Funa L = Funa latisinuata, Vexitomina G = 
Vexitomina garrardi, Pilsbryspira N = Pilsbryspira nympha, Aforia A = Aforia abyssalis, Lophiotoma L = Lophiotoma leucotropis , 
Polystira A = Polystira albida, Micantapex P = Micantapex parengonius, Borsonia O = Borsonia ochraea, Ophiodermella I = 
Ophiodermella inermis, Oenopota L = Oenopota levidensis, Tropidoturris F = Tropidoturris fossata, Anarithma M = Anarithma metula, 
Glyphostoma C = Glyphostoma Candida, Genota N = Genota nicklesi, Benthofascis = Benthofascis biconica, Conus V = Conus 
ventricosus, Tomopleura V = Tomopleura reevei, Eucithara S = Eucithara stromboides, Mangelia N = Mangelia nebula, Mangelia P = 
Mangelia powisiana, Thatcheria M = Thatcheria mirabilis, Philbertia P = Philbertia purpurea, Philbertia L = Philbertia linearis, Daphnella 
R = Daphnella reeveana, Gymnobela E = Gymnobela emertoni, Teretiopsis L = Teretiopsis levicarinatus, Abyssobela A = Abyssobela 
atoxica, Taranis M = Taranis moerchi. 



ribbon and loss of the odontophore. 

Clade 19 is made up of various taxa formerly included in 
the Borsoniinae and Clathurelhnae with the addition of 
Oenopota (Oenopotinae). The apomorphies defining the 
nodes are very unsatisfactory with many reversals. More 
characters need to be analysed in these taxa to achieve better 
resolution. 

Borsonia and Ophiodermella (Node 21) have posteriorly 
situated rhynchodeal sphincters, and fusiform shells. The 



taxa in the other clade (Node 22) have extensible buccal lips 
and hollow radular teeth with large bases. Although the 
Oenopotinae have been previously thought to have close 
affinities with the Mangehinae, they do have acinous salivary 
glands, rather than the tubular type associated with the latter 
subfamily. 

A clade comprising Anarithma and Glyphostoma is defined 
(Node 24) by three characters; a posteriorly situated rhyn- 
chodeal sphincter, a single salivary duct and apertural orna- 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



155 



ment. Glyphostoma has long slender radular teeth and has 
been separated in the family Clathurellinae (McLean, 1971). 
Anarithma has been classified in the Diptychomitrinae (= 
Mitrolumninae), but Kilburn (1986) could see no significant 
differences from the Borsoniinae. 

Taxa normally classified in the Borsoniinae (Ophioder- 
mella, Borsonia, Tomopleura, Micantapex, Tropidoturris and 
Anarithma) do not form a monophyletic group in any of our 
analyses. For this reason, in the classification derived form 
this study we are leaving these taxa, along with Glyphostoma 
and others in informal groupings within the subfamily 
Clathurellinae. 

Benthofascis (Conorbinae) and Conus (Coninae) (Node 
27) share a number of characters. They lack an anterior 
sphincter to the buccal tube, but have have an intermediate 
sphincter instead. Both have accessory salivary glands and 
retain an operculum. Additionally, both genera show resorp- 
tion of the inner shell whorls. Although Genota (Node 26) is 
usually classified in the Conorbinae, it lacks an operculum. 

Taxa from Node 28 onwards have tubular salivary glands 
and most have sculptured protoconchs. The Mangeliinae 
(Node 30), represented by Eucithara and Mangelia, are a 
well-defined group with the distinctive buccal tube introvert, 
and protrusive lips of the buccal tube. Taxa from Node 32 
have a muscular bulb made up of only one muscle layer and 
lacking the connective tissue layer, with additionally, an anal 
sinus located at the suture. Thatcheria (Node 32) has many 
characters in common with the Daphnellinae and until many 
more daphnellines have been examined anatomically it can 
be classified with them. However a great range of foregut 
anatomy is found in the Daphnellinae and it may be that the 
group is paraphyletic. At the extreme end of the tree (Node 
36) are taxa which have lost many foregut characters such as 
radula, proboscis and glands. Taranis has been classified in a 
separate subfamily Taraninae (Kantor & Sysoev, 1989), but 
it has so few characters that its relationships are obscure. It 
may be a highly derived daphnelline. 

Conclusions 

Our studies have shown that several major autapomorphies 
associated with the Conoidea have developed independently 
in separate clades. Also there has been parallel loss of foregut 
structures. Some of the more important of these are briefly 
discussed below. 

Hollow, enrolled 'hypodermic style' radular teeth are con- 
sidered a distinctive feature of the conoidean feeding mecha- 
nism. Our analysis shows that hollow teeth have been 
independently derived at least five times in the evolution of 
the Conoidea. In Imaclava the hollow marginal teeth seem to 
have developed from the enrolling of the flattened drilliine- 
type of marginal teeth. In Toxiclionella, the hollow teeth 
were derived from wishbone teeth similar to those of Clav- 
atula or maybe from solid teeth like those of Pseudome- 
latoma. Hollow teeth are found in many Terebridae and are 
thought to have been derived from solid teeth via semi- 
enrolled intermediate forms such as found in Hastida bacillus. 
The enrolled teeth of Pilsbryspira (Zonulispirinae) may have 
been derived by enrolling of the crassispirine type of wish- 
bone tooth. The hollow teeth of the higher conoideans such 
Clathurellinae, Coninae, Mangeliinae and Daphnellinae in 
all their various forms may represent another separate deriva- 
tion. The radular caecum found in some Terebridae was 
derived independently of that found in the higher turrids 



(Clathurellinae, Oenopotinae, Mangeliinae, Daphnellinae) 
and Coninae. 

The rhynchodeal introvert found in some Daphnellinae, is 
also found in all Terebridae (including pervicaciines). If our 
ideas concerning the relationships of the Terebridae are 
correct, then the structure was evolved independently in the 
two groups. 

A buccal mass situated at the base of the proboscis is 
considered to be a diagnostic character of the Conoidea 
(Ponder, 1973). However, in Turricula nelliae the buccal 
mass was shown to be located at the distal end of the 
proboscis (Taylor, 1985; Miller, 1990). We now know that a 
distally-shifted buccal mass seems to be common feature of 
the Clavatulinae and is found also in Pilsbryspira 
(Zonulispirinae) and Strictispira (Strictispirinae) which lacks 
the venom apparatus. 

One surprising trend seen in at least four clades is the loss 
of the venom apparatus. In the Daphnellinae, Taraninae and 
some Terebrinae this is associated with the loss of the 
proboscis and radular apparatus. Pervicaciinae have a well 
developed radula apparatus but no proboscis or venom gland. 
By contrast, Strictispira which also lacks the venom gland, has 
a proboscis, a distally-located buccal mass and a robust radula 
apparatus. 



Relationships and status of Terebrinae and 
Pervicaciinae 

Some controversy concerns the status of the Terebrinae and 
Pervicaciinae. Rudman (1969) and Taylor (1990) suggested 
an independent origin for the two groups. However, anatomi- 
cal studies of more species is revealing some shared apomor- 
phies which suggest a common origin. 

Although both subfamilies possess elongate multi-whorled 
shells there are large anatomical differences between the two 
groups. The family Pervicaciidae was orginally proposed by 
Rudman (1969) for Pervicacia tristis, a terebriform species 
with no proboscis and venom apparatus, but with an odonto- 
phore and a radula with a strong membrane and two sickle- 
shaped, solid teeth in each row. It is now known, that many 
more 'terebrids' (Duplicaria species and others) share these 
characters and should be included in the family (Taylor, 
1990). Other characters of pervicaciids include a rhynchodeal 
introvert and a septum in some species. 

Most of the radulate Terebrinae s.s. possess hollow and 
barbed, radular teeth, similar to those seen in Conus and the 
Clathurellinae. However, some Hastula species possess an 
odontophore and Hastula bacillus has partially-solid teeth 
(Taylor & Miller, 1989). This discovery demonstrates that the 
Terebridae must be derived from a lower conoidean with an 
odontophore and radular ribbon, rather than from some 
group such as the Clathurellinae, which have lost these 
structures. 

The accessory proboscis structure is an unusual organ 
found in some Terebrinae, and is known from Hastula 
bacillus, H. aciculina, H. imitatrix, H. raphanula, Terebra 
affinis and T. pertusa (Miller 1971, Taylor , 1990; Auffenberg 
& Lee, 1988; Taylor, unpub.). Some terebrines, for example 
Terebra subulata, also possess a septum dividing the rhyn- 
chocoel (Miller, 1971; Taylor 1990). We have found an 
accessory proboscis structure in the western Australian spe- 
cies Duplicaria kieneri, and Duplicaria colorata (recently 
described as a Hastula by Bratcher (1988)), which otherwise 



156 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



have an anatomy similar to Pervicacia. 

Although the pervicaciines and terebrines apparently differ 
considerably in foregut anatomy, they share a a number of 
characters which suggest a common origin (Table 5). The 
idea that the Terebrinae and Pervicaciinae were derived 
separately (Rudman, 1969; Taylor, 1990) is rejected. Charac- 
ters in common between the two groups are: the elongate 
multi-whorled shell, the rhynchodeal introvert, and in some 
species the rhynchodeal septum and accessory proboscis 
structure. Thus, we propose that the common ancestor of the 
combined Pervicaciinae and Terebrinae clade would have 
possessed a rhynchodeal introvert, a proboscis, an odonto- 
phore, a radula with two solid, sickle-shaped, marginal teeth 
in each row, a venom gland, a pair of acinous salivary glands, 
a pair of accessory salivary glands, an accessory proboscis 
structure and a rhynchodeal septum. 

Species in the Pervicaciinae clade have lost the proboscis, 
venom gland and accessory salivary glands. In the Terebrinae 
clade, the solid radular teeth were transformed into semi- 
enrolled and then hollow teeth. The odontophore was also 
progressively lost. Species with hollow teeth have developed 
a radular caecum. Other, more-derived terebrines and possi- 
bly pervicaciines, have lost virtually all the foregut structures, 
with the rhynchodeal introvert becoming the main feeding 
organ (Taylor, 1990). 

Because the radula with solid, sickle-shaped marginal teeth 
and well developed odontophore, is regarded as one of the 
least-derived for the Conoidea, we regard the Pervicaciinae/ 
Terebrinae clade as an early branch from the rest of the 
Conoidea. If our hypothesis of relationships is correct, then 
the hollow, barbed teeth, the radular caecum, the rhyn- 
chodeal introvert, and rhynchodeal septum of the terebrids, 
have been derived independently of those similar structures 
found in the Daphnellinae and Clathurellinae. 

Status of Conidae 

Despite the distinctive shell form and high species diversity of 
the group, we have little anatomical evidence to support the 
separation of Conus at family-level from other higher turrids. 
We propose only sub-family status for the group. Every 
anatomical character-state of the conine foregut is shared 
with species of Clathurellinae and Conorbinae. Some Conus 
species possess a snout gland in the rhynchocoel, but this 



organ has been little studied. Conus species also have a 
distinctive osphradium with the multidigitate leaflets (Taylor 
& Miller, 1989). However, the detailed structure of the 
osphradium has been studied in only a few species of Tur- 
ridae, but at least in some Aforia species (Cochlespirinae) 
there are similar digitate osphradial leaflets (Sysoev & Kan- 
tor, 1988). The resorption of the inner shell whorls has been 
used as a diagnostic character of conines (Kohn, 1990), but 
the occurrence of this feature has been little studied in other 
conoideans, although it is present in Benthofascis (Conorbi- 
nae). 



CLASSIFICATION OF CONOIDEA 



Introduction 

Although many of the subfamilial names (as well as apparent 
synonyms) currently-used within the Turridae were intro- 
duced in the 19th or early 20th century, no detailed and 
well-documented classification was developed in these earlier 
works. Most authors based their classifications exclusively on 
shell characters, although Stimpson (1865) used radula data 
and Fischer (1887) divided the Conoidea into four subfamilies 
solely by opercular characters. The rather detailed classifica- 
tion of Casey (1904) who recognised eight tribes within the 
Turridae (Donovaniini are not conoideans), was based on 
both opercular and shell characters. 

Thiele (1931) classified turrids into three subfamilies con- 
tained within the family Conidae, with the Terebridae as a 
separate family. Diagnoses of the turrid subfamilies mainly 
consisted of combinations of such characters as 'opercu- 
late-inoperculate' and 'toxoglossate-nontoxoglossate denti- 
tion'. This was the first classification where the taxonomic 
difference between toxoglossate and nontoxoglossate radulae 
was definitely indicated. An elaboration of this classification 
was developed by Wenz (1938) who recognised five subfami- 
lies of Turridae as well as the Conidae and Terebridae. 

The classification of Powell (1942, 1966) provided a great 
stimulus to conoidean taxonomy, and is used, with modi- 
fications, by almost all authors concerned with Turridae. 
Powell recognized nine subfamilies which were based prima- 
rily on shell characters, although radular and opercular 



Table 5. Comparison of character states between Pervicaciinae and Terebrinae. 



Character 



Pervicaciinae 



Terebrinae 



Shell shape 
Radular teeth 
Odontophore 
Radular caecum 
Venom gland 

Proboscis 
Salivary glands 
Accessory salivary glands 
Rhynchodeal introvert 
Rhynchodeal septum 
Accessory proboscis structure 
Eyes 
Operculum 



Multiwhorled 

Solid sickle-shaped 

Present 

Absent 

Absent 

Absent 
Present 
Absent 
Present 

Present in some 
Present in some 
Absent in all? 
Present 



Multiwhorled 

If present, usually hollow enrolled marginals 

Present in some Hastula species 

Present in hollow-toothed forms 

Present in all with radula & proboscis 

absent in others 

Present in all radulate forms 

Present in many species 

Present in some species 

Present 

Present in some 

Present in some 

Present 

Present 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



157 



features were also used. Powell believed that the hypodermic 
toxoglossate dentition could develop independently in differ- 
ent lineages and, more importantly, that the appearance of 
toxoglossate radula was not a significant reason for separating 
groups at the subfamilial level. As a result, he classified some 
taxa having quite different radular types (including both solid 
and hollow marginal teeth) within a single subfamily. 

Morrison (1966) followed Thiele in recognizing a funda- 
mental difference between groups with solid (= nontoxoglos- 
sate) and hollow (= toxoglossate) marginal teeth. He 
suggested a separation at the family level using the families 
Turridae (with subfamilies Drilliinae, Clavatulinae and 
'Lophiotominae or Crassispirinae'), Mangeliidae and 
Tseudomelatominae'. 

The subfamily classification of Powell was considerably 
revised by McLean (1971), who adhered strictly to the 
principle of grouping together genera with the same type of 
radula. He also added six subfamilies to Powell's classifica- 
tion; three of these being described as new (Clathurellinae 
H. & A. Adams, erroneously). Several subfamilies were 
recognised (or retained after Powell) on shell characters, 
but which share the same radular type, and some of these 
seem to be rather poorly documented. However, McLean's 
classification which includes 15 subfamilies is at present the 
most detailed and well developed. 

In a continuing series of papers concerning South African 
Turridae, Kilburn (1983, 1985, 1986, 1988), adopted a prag- 
matic approach (Kilburn, 1983 p. 550 ' . . . any practical 
subdivision is better than none ...'), and revised to some 
extent the composition of subfamilies which he studied. He 
also synonymized the Diptychomitrinae (= Mitrolumninae = 
Mitromorphinae) with the Borsoniinae. 

Bogdanov (1986, 1987, 1990) described a new subfamily 
Oenopotinae separating the operculate Oenopota and its 
relatives from the Mangeliinae. Additionally, the subfam- 
ily Taraninae was recently re-instated (Kantor & Sysoev, 
1989). 

Some nomenclatural changes in the names and authorships 
of several subfamilies were made by Cernohorsky (1972, 
1985, 1987), and Ponder and Waren (1988). 

A different viewpoint was taken by Bouchet and Waren 
(1980) in their study of North Atlantic deep-sea Turridae. 
They avoided the use of any subfamilial divisions, considering 
the present classification of Turridae to be artificial and based 
mainly on (p. 5) ' . . . more or less randomly selected shell 
characters'. 

At present there is no completely agreed classification of 
Turridae, nor is there any agreement on which are the 
taxonomically important characters. The existing variants of 
turrid classification are based almost exclusively on shell, 
radular and opercular features. 

The Terebridae have similarly been classified mainly on 
shell characters. H. & A. Adams (1853) and Cossmann 
(1896) divided the Terebridae into two subfamilies, includ- 
ing the Pusionellinae as the second subfamily. Pusionella is 
now known to belong to the turrid subfamily Clavatulinae. 
A separate family, the Pervicaciidae, was proposed by 
Rudman (1969) for Pervicacia tristis. However, Bratcher & 
Cernohorsky (1987) included Pervicacia and similar forms 
in the Terebridae. Taylor (1990) confirmed the distinctive- 
ness of Pervicacia, and showed that many other terebrids 
should be included in the family Pervicaciidae. 

The Conidae have long been considered as a fairly homo- 
geneous group, the main problems have concerned the limits 



of the family and whether taxa such as Cryptoconus, Conor- 
bis and Genota should be included. Cossmann (1896) for 
example, included them in the subfamily Conorbinae within 
the Conidae, whilst Powell (1966) includes this subfamily in 
the Turridae. 



New classification proposed 

As a result of our analysis of foregut characters throughout all 
the conoidean higher taxa we propose a new classification of 
the superfamily. This classification represents a rather con- 
servative compromise position. Although in principle the 
classification should be based upon the results of the phyloge- 
netic analysis, we were constrained by the rather poor 
resolution obtained with our data set. Moreover, only a 
rather small subset of conoidean species have been examined 
in any detail. Information from taxa not included in the 
cladistic analysis (mainly radular characters) has also been 
used in constructing the classification. An example of the 
problem is the family Turridae, which comprises the four 
subfamilies with wishbone marginal teeth, plus the 
Zonulispirinae. The cladistic analysis suggests two different 
clades for these subfamilies. This is certainly possible, but the 
branches are supported by rather few, and perhaps weak 
apomorphies. Despite the deficiencies this is the first compre- 
hensive classification of the Conoidea which includes ana- 
tomical characters. Below we give descriptions of shell, 
radula and foregut characters for each of the higher taxa that 
we recognise. Some of the taxa have only provisional status. 
For example, the subfamily Clathurellinae has been divided 
up into five informal groups; it may well be polyphyletic, but 
we have insufficient evidence to resolve the situation. Simi- 
larly, we are uncertain of the status of the Conorbinae and 
Taraninae. 



Summary of proposed classification 

Superfamily Conoidea 

Family Drilliidae (ICZN pending) 
Family Terebridae 

Subfamily Pervicaciinae 
Terebrinae 
Family Pseudomelatomidae 

Family Strictispiridae 

Family Turridae 

Subfamily Clavatulinae 
Crassispirinae 
Zonulispirinae 
Cochlespirinae 
Turrinae 

Family Conidae 
Subfamily Clathurellinae 
Coninae 
Conorbinae ? 
Oenopotinae 
Mangeliinae 
Daphnellinae 
Taraninae ? 



158 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



DIAGNOSES OF HIGHER TAXA 

Family Drilliidae Morrison, 1966 (ICZN pending) 

Shell of small to medium size (usually 15-25 mm, up to 50 
mm), claviform (with a more or less high spire, and a 
relatively short, truncated base). Anterior canal indistinct, 
short or moderately elongate. Anal sinus on the shoulder, 
rather deep, often sub-tubular when a parietal tubercle is 
present. Sculpture usually well developed. Protoconch pauci- 
or multispiral, smooth or, sometimes, carinate (from the 
second whorl or, rarely, from the beginning). Operculum 
with terminal nucleus. 

Radula. With strong radular membrane, five teeth in each 
row, with in some species the complete loss of the central 
tooth and reduction of the laterals. Rachidian tooth small, 
with a prominent central cusp and, often, smaller lateral 
denticles. Lateral teeth are typically broad and curved, 
comb-like, with many small cusps the outermost being 
smaller. Marginal teeth have a variable morphology from 
simple and flat, sometimes with a weak accessory limb, to 
enrolled. In at least one species (Imaclava unimaculata) , 
marginal teeth are hollow and enrolled, whilst the radula as a 
whole is similar to that of other drilliids. 

Foregut. Proboscis moderately long, with one or two distal 
sphincters and sometimes a mid-buccal tube sphincter. Buccal 
mass at base of proboscis, odontophore well-developed, 
cartilages either separated or fused. Two acinous salivary 
glands with two ducts. No accessory salivary glands. Venom 
gland with uniform histology along its length. Retractor 
muscle of the radular sac passes through the nerve ring and 
joins the columellar muscle. 

Remarks. The anatomy and radula are known for only a 
very few species of Drilliidae. This prevents us from introduc- 
ing any subfamilial classification of this possibly complex 
family. 



Family Terebridae Morch, 1852 

Elongate, multiwhorled shells, with small quadrate to trian- 
gular apertures. Siphonal canal short. Anal sinus not visible. 
Shell ornament of low axial ribs and grooves, spiral grooves, 
a few species with tubercles, shells often smooth and pol- 
ished. Protoconch of 1.5-5 whorls. Operculum rounded with 
terminal nucleus. Radula with solid sickle-shaped teeth, 
hollow harpon-like teeth or absent. Rhynchodeal introvert 
present. Accessory proboscis structure and rhynchodeal sep- 
tum present in some species. Proboscis present or absent. 
Odontophore present in some species. Radular caecum 
present in some. Acinous salivary glands present. Accessory 
salivary glands present in some species. Venom gland present 
or absent. 



Subfamily Pervicaciinae Rudman, 1969 

Shells medium to large, elongate, multiwhorled, anterior 
canal short, ornament low axial ribs, spiral grooves, often 
with a subsutural groove. Aperture quadrate. Operculum 
rounded with terminal nucleus. 

Radula. With strong radula ribbon, two rows of sickle- 
shaped solid marginal teeth. 



Foregut. Rhynchodeal introvert. Rhynchodeal septum and 
accessory proboscis structure present in some species. Pro- 
boscis absent. Extensible buccal lips present in some species. 
Odontophore with two cartilages. Two acinous salivary 
glands and ducts. Venom gland and accessory salivary glands 
absent. 



Subfamily Terebrinae Morch, 1852 

Shells medium to large, elongate, multiwhorled. Small quad- 
rate to triangular aperture. Short siphonal canal. Shells often 
smooth and polished. Shell ornament of low axial and spiral 
ribs and grooves. 

Radula. Where present, long, hollow marginal teeth with 
narrow bases, barbed or unbarbed. Hastula bacillus has 
semi-enrolled teeth with a distal solid blade. Many species 
have no radula. 

Foregut. Rhynchodeal introvert present. Rhynchodeal sep- 
tum and accessory proboscis structure present in some spe- 
cies. Proboscis long, medium or absent. Odontophore with 
cartilages present in some Hastula species. Radula caecum 
present in many radulate species. Acinous salivary glands 
with two ducts usually present. Accessory salivary glands 
present in some species. Venom gland present or absent in 
radula-less species. 



Family Pseudomelatomidae Morrison, 1966 

Shells of medium to large size (35-77 mm), fusiform. Ante- 
rior canal moderately elongate. Anal sinus on the shoulder. 
Protoconch smooth. Operculum with terminal or subcentral 
nucleus. Egg capsules dome-shaped, with an operculum. 

Radula. With strong radular membrane; three teeth in each 
radular row. Rachidian is large and rectangular with a large, 
curved and pointed, central cusp and smaller lateral cusps. 
Marginal teeth are solid, simple and curved. 

Foregut. Proboscis very long, no anterior buccal tube 
sphincter; buccal mass basal or posterior of the proboscis 
base. Oesophagus elongated between the buccal mass and 
nerve ring in Pseudomelatoma . Odontophore very large with 
fused cartilages. Acinous salivary glands, paired in 
Pseudomelatoma, but unpaired with a single duct in Hormo- 
spira. No accessory salivary glands. Venom gland with uni- 
form histology. 



Family Strictispiridae McLean, 1971 

Shell of rather small size (usually 15-20 mm), claviform. 
Anterior canal short or indistinct. Sculpture well developed. 
Deep subtubular sinus is situated on the concave shoulder 
and bordered with well developed parietal callus. Protoconch 
smooth, multispiral. Operculum with terminal nucleus. 

Radula. with strong radular membrane; 2 teeth in each 
row, central and lateral teeth absent (latter maybe diapha- 
nously on optical preparations). Marginal teeth solid, awl- 
shaped, with pointed tips, a broad base and a mid-tooth 
flange . 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



159 



Foregut. Proboscis short; buccal mass located near the 
proboscis tip, odontophore very large and muscular with 
separate cartilages. Acinous salivary glands small and paired, 
no accessory salivary gland, no venom apparatus. 

Remarks. This small family possesses unique radular teeth 
and anatomy, but study of further material is necessary. 



elongated oesophagus between the buccal mass and nerve 
ring). Moreover, it is also similar to clavatulines in shell 
characters and in its operculum with mediolateral nucleus. 
On the other hand, Turricula differs in both shell and 
anatomical characters from those of other 'Turriculinae'. 
Thus we transfer this genus, as well as Makiyamaia which has 
similar characters, to the subfamily Clavatulinae. 



Family Turridae H. & A. Adams, 1853 

Radula always with a membrane with either 3 radular teeth in 
a row (central being small or weak), 4 (central lost, laterals 
diaphanous) or with only marginals. Marginal teeth usually 
wishbone type, rarely enrolled and hollow. Odontophore 
always present. Radular sac not subdivided into short and 
long arms. Venom gland always present. Salivary glands 
always acinous. Accessory salivary gland either present or 
absent. Operculum present. 



Subfamily Clavatulinae Gray, 1853 

Shell medium-sized (usually 15-30 mm, maximum 60 mm), 
variable in form. Anterior canal moderately long, sometimes 
short or trun-cated. Whorls usually adpressed below the 
suture. Anal sinus located on the shoulder slope, rather deep 
but sometimes indistinct. Protoconch smooth, of 1.5-3 
whorls. Axial sculpture predominates or the sculpture is 
subobsolete and the shell surface is glossy. Operculum ovate, 
with medio-lateral nucleus. Egg capsules lens-shaped, verti- 
cally orientated, without an operculum. Capsules attached to 
the substratum by a stalk on the edge. 

Radula. Strong radular membrane with 3 to 2 teeth in each 
row. Central tooth with large, very thin, inconspicuous, basal 
plate and centrally thickened area with a single cusp. Central 
tooth sometimes absent (Toxiclionella s.s.). Lateral teeth 
absent. Marginal teeth usually robust wishbone type; hollow 
harpoon-shaped and barbed in Toxiclionella. 

FOREGUT. Epithelium of posterior rhynchocoel not glandu- 
lar and continuous with proboscis. Moderately long proboscis 
with 1 or 2 anterior buccal tube sphincters. Protrusive lips of 
the buccal tube may be present (Turricula). Buccal mass 
distal except Clavatula diadema in which it is basal but lies 
within the proboscis. Odontophore medium to small in size, 
cartilages unfused (except in Toxiclionella). Salivary glands 
acinous, usually paired. Single salivary duct in Clavatula 
caerulea. Single accessory salivary gland in Toxiclionella. 
Anterior venom gland ciliated. Oesophagus elongated 
between buccal mass and nerve ring. 

Remarks. Some species in this subfamily possess hollow 
'toxoglossate' radular teeth associated with strong radular 
membrane, sometimes, with central teeth. The anatomy and 
conchological characters of 'toxoglossate' clavatulines are, 
however, quite similar to those of "nontoxoglossate' ones. 
Thus at present we do not consider the appearance of 
hollow teeth in Toxiclionella to be a taxonomic character of 
subfamilial importance and therefore follow Kilburn (1986) 
in classifying Toxiclionella with other clavatulines. 

The genus Turricula Schumacher, 1817 appears very simi- 
lar to clavatulines in both radular characters and anatomy 
(the distal buccal mass, ciliated anterior venom gland, 



Subfamily Crassispirinae Morrison, 1966 

Shell of medium to small size (usually 10-20 mm, sometimes 
up to 70 mm), claviform to fusiform. Anterior canal usually 
short. Anal sinus on the whorl shoulder, parietal callus above 
the sinus often well developed. Spiral and axial sculpture 
often strong. Protoconch usually paucispiral, initially smooth, 
later sometimes with axial (rarely spiral) folds. Operculum 
with terminal nucleus. 

Radula. Strong radular membrane and 4, 3 or 2 teeth in 
each row. Central tooth when present (Turridrupa) is thin, 
quadrate and unicuspate, lateral teeth usually absent but 
weak and vestigial in Crassispira and Crassiclava. Marginal 
teeth, robust wishbone type or long flat teeth with a slender 
accessory limb. Ptychobela has hollow teeth formed from two 
components. 

FOREGUT. Proboscis moderately long with two anterior buc- 
cal tube sphincters. Epithelium of posterior rhynchocoel 
continuous with proboscis (Funa latisinuata). Buccal mass 
situated at the proboscis base in its contracted state. Odonto- 
phore medium to small, with fused cartilages. Salivary glands 
acinous, fused, ducts paired. Anterior venom gland ciliated in 
some species. Oesophagus elongated behind buccal mass in 
some species. 

Remarks. This most large and diverse subfamily of Turridae 
is defined chiefly on shell and radular characters (i.e. rather 
small claviform shells with wishbone radular teeth). Data on 
the anatomy of its representatives are still unsufficient to 
decide certainly whether the subfamily is of mono- or poly- 
phyletic origin. 



Subfamily Cochlespirinae Powell, 1942 

Shell of medium to large size (usually 20-40 mm, up to 100 
mm), narrow to broadly fusiform or pagodiform. Anterior 
canal moderately elongate, rarely short or very long. Sculp- 
ture variously developed, often with smooth shoulder, and 
usually with rather short axial ribs below the shoulder, and 
spiral riblets. Anal sinus usually deep, situated on the shoul- 
der (sometimes on its lower part). Protoconch usually multi- 
spiral, smooth or, sometimes, initially smooth and carinated 
or spirally or axially lirate on subsequent whorls. Operculum 
with terminal nucleus. 

Radula. Strong radular membrane, with three, four? (see 
discussion of radula p. 135) or two teeth in each row. Central 
tooth weak, unicuspid or absent. Marginal teeth of robust 
wishbone type. 

Foregut. Proboscis usualy long, with one or two anterior 
buccal tube sphincters. Buccal mass basal, muscular buccal 
lips may be present or absent. Odontophore small, cartilages 
4, 2 or absent, fused or separate. Salivary glands acinous. 



160 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



paired or fused. Single accessory salivary gland in Aforia. 

Remarks. Since the type-genus of the subfamily Turriculi- 
nae, Turricula Schumacher, 1817, is transferred to the Clav- 
atulinae (see above), the next available name for this group is 
Cochlespirinae Powell, 1942. 



Subfamily Zonulispirinae McLean, 1971 

Shells rather small (15-25 mm), claviform. Anterior canal 
usually short, sometimes moderately long. Predominantly 
spiral scuplture, well developed. Protoconch multispiral, ini- 
tially with smooth whorls, then with oblique axial riblets. 
Anal sinus on the shoulder, often sub-tubular, with well 
developed parietal callus. Operculum with terminal nucleus. 

Radula. With strong membrane and marginal teeth in each 
row. Teeth semi-enrolled, to rolled, hollow teeth with narrow 
base. Tips may be barbed or unbarbed. 

Foregut. Proboscis long, with a single distal buccal tube 
sphincter. Buccal mass distal. Odontophore small with two 
unfused cartilages. Buccal lips present. Salivary glands fused. 
Anterior of venom gland ciliated. Oesophagus elongated 
between the buccal mass and nerve ring. 



Subfamily Turrinae H. & A. Adams, 1853 

Shell usually of medium to large size (up to 110 mm), 
fusiform. Anterior canal elongated and narrow, rarely trun- 
cated. Anal sinus on the whorl periphery. Axial sculpture 
weak or absent. Protoconch smooth in its initial part, subse- 
quent whorls axially costate; paucispiral protoconchs smooth. 
Operculum with terminal nucleus. Egg capsules dome- 
shaped, operculate. 

Radula. Strong radular membrane, 2-3 teeth in each row. 
Central tooth either well-developed, small or absent, quad- 
rate to rectangular with a strong central cusp. Lateral teeth 
absent. Marginal teeth of robust wishbone type. 

Foregut. Proboscis moderately long, rhynchostomal 
sphincter posterior, a single distal buccal tube sphincter, 
protrusive lips of buccal tube present. Buccal mass basal. 
Odontophore small with fused cartilages. Salivary glands 
paired. No accessory salivary glands. Anterior part of venom 
gland ciliated. 



Family Conidae Fleming, 1822 

Radula consisting of hollow marginal teeth only. Radular 
membrane absent. Radular diverticulum divided into short 
and long arms. Odontophore absent. Radula and venom 
gland may be absent. Salivary glands acinous or tubular. 
Accessory salivary gland either present or absent. Operculum 
either present or absent. 



Subfamily Clathurellinae H. & A. Adams, 1858 

Shell small to rather large, fusiform to biconic. Anterior canal 
short or indistinct to moderately elongate. Sculpture pre- 
dominantly spiral in most genera. Anal sinus deep to very 



shallow, on the shoulder slope or on the periphery. Col- 
umella with or without pleats. Protoconch usually paucispiral, 
smooth, sometimes carinate or weakly spirally ribbed, rarely 
axially costate on its last whorl. Operculum with terminal 
nucleus present, vestigial or absent. 

Radula. Awl- or harpoon-shaped marginal teeth, without 
(very rarely with) solid base, tooth cavity opens terminally at 
the proximal end in vast majority of species. 

Foregut. Proboscis short to long, 1 or 2 anterior buccal 
tube sphincters, buccal mass basal. Short buccal lips in 
Tropidoturris. Odontophore absent, radular caecum present 
— divided by septum in Bathytoma (Micantapex) . Salivary 
glands tubular in Borsonia, acinous in others, paired, single 
or absent. Single accessory salivary gland present in some 
species. Venom gland with uniform histology. No elongation 
of oesophagus. 

Remarks. This subfamily comprises species classified by 
other workers in the subfamilies Borsoninae and Clathurelli- 
nae. Being very variable in both anatomical and shell charac- 
ters, the subfamily may be of polyphyletic origin. More 
species need to be studied anatomically before any satisfac- 
tory classification can be attempted. The subfamily is defined 
mainly by the character of the radular teeth. Several groups 
of genera can be isolated within Clathurellinae according to 
shell characters. 

'Clathurellid' group is characterized by medium-sized 
shells (usually 10-25, up to 40 mm), with a moderately 
elongate siphonal canal, and a well developed, often cancel- 
late sculpture. Columella without pleats, but both inner and 
outer lips may be denticulated; anal sinus deep located on the 
shoulder. Protoconch usually multispiral, last whorls with a 
pronounced medial carination and, sometimes, weak axial 
lamellae on the lower half. A distinctive feature of this group 
is densely granulated shell surface of most genera (except of 
one subgenus of Glyphostoma and, probably, Nannodiella). 
Operculum absent. Radular teeth long and slender, slightly 
curved, without a solid base. 

'Bathytomid' group. Shell of medium to rather large size 
(usually 20-30, up to 70 mm), more or less biconic. Sculpture 
usually well developed, entirely spiral, ribs often gemmulated 
by growth lines; typically there is a peripheral tuberculated 
flange. Anal sinus rather deep, located on the whorl periph- 
ery. Columellar pleats strong to obsolete. Protoconch of 
1.5-3 whorls, smooth or minutely papillated. Operculum with 
terminal nucleus. Radular teeth either long, with more or less 
terminal opening, or short, with large cylindrical solid base 
and lateral opening. 

'Borsoniid' group. Shell of rather small to medium size 
(usually 15-25, up to 62 mm), fusiform. Anterior canal 
moderately elongate, sometimes long. Both spiral and axial 
sculpture may be present. Columellar pleats weak or absent. 
Anal sinus on the shoulder slope. Protoconch of 1-2 smooth 
whorls. Operculum fully developed, small or absent. Radular 
teeth long, without solid base, open terminally, or, rarely, 
short, with large cylindrical base, open laterally. Egg capsules 
dome-shaped, with an operculum. 

'Mitromorphid' group. Shell small (usually 4-8, up to 17 
mm), biconic and 'mitriform'. Anterior canal very short or 
indistinct. Aperture narrow, columella with or without teeth, 
outer lip usually denticulated, anal sinus shallow and subsu- 
tural. Sculpture predominantly or entirely spiral. Protoconch 
of 1.5-2 smooth whorls. Operculum absent. Radular teeth 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



161 



rather short, of 'candle flame' shape, open terminally. 

'Tomopleurid' group. Shell rather small to medium sized 
(6-7 to 37 mm), claviform, with flattened whorls. Anterior 
canal short. Anal sinus on the shoulder or just below it, 
moderately deep. Columellar pleats absent. Sculpture 
entirely spiral (except often raised growth lines), consisting of 
well developed ribs or heavy keels. Protoconch pauci- or 
multispiral. In the former case it is smooth or with minute 
spiral striae or papillae, sometimes carinated; in the latter 
case first 1-3 whorls with the same sculpture, later ones with 
axial ribs and, sometimes, minute spiral striae. Operculum 
with terminal or eccentric nucleus, sometimes absent. Radu- 
lar teeth short or long and slender, without solid base, open 
terminally. 



Subfamily Conorbinae De Gregorio, 1890 

Shell of medium size (up to 40 mm), biconic. Anterior canal 
short, aperture long and narrow. Sculpture entirely spiral 
except the growth lines. Anal sinus on the shoulder or almost 
sutural, relatively deep. Protoconch multispiral, smooth or 
spirally striated on later whorls. Operculum present or, 
absent in Conorbis. 

Radula. Hollow, marginal teeth with barbed tips and nar- 
row bases (Conorbis, Thiele, 192 fig 460; Benthofascis, 
Powell, 1966, fig. 125). 

FOREGUT. These observations are based on Benthofascis. 
Rhynchostomal sphincter posteriorly situated. Proboscis 
moderately long, not folded telescopically as in Conus. Distal 
sphincter of buccal tube absent, intermediate sphincter 
present. Middle part of buccal tube lined with glandular 
epithelium. Single acinous salivary gland with two ducts. 
Single accessory salivary gland. Venom gland with uniform 
histology, muscular bulb with two muscular layers. No snout 
gland. 

Remarks. The status of this subfamily is uncertain due to 
lack of any anatomical information on Conorbis. We have 
excluded Genota on the basis of shell morphology and the 
absence of the operculum. 



Snout gland present in many species. Distal buccal tube 
sphincter absent, intermediate sphincter present. Middle part 
of buccal tube is lined with glandular epithelium. Buccal mass 
basal. Single acinous salivary gland with one or two ducts. 
Single accessory salivary gland. Venom gland of uniform 
histology, muscular bulb often with many muscular layers. 



Subfamily Oenopotinae Bogdanov, 1987 

Shell of small to medium size (usually 10-15, up to 30 mm), 
oval to fusiform. Anterior canal rather short. Both spiral and 
axial sculpture well developed. Anal sinus on the shoulder, 
shallow, and often indistinct. Protoconch paucispiral, pre- 
dominantly (sometimes entirely) spirally sculptured. Opercu- 
lum with terminal nucleus present, vestigial, or rarely absent. 
Egg capsules dome-shaped, with an operculum. 

Radula. Radular teeth with rounded or cylindrical solid 
base and hollow shaft, sometimes with barbed tip; rarely 
teeth vestigial; tooth cavity opens laterally between the shaft 
and the base. 

Foregut. Proboscis either long, or short and folded in 
contracted state. Distal sphincter present or absent. Buccal 
lips large, may be inverted into the buccal cavity. Buccal mass 
basal. Salivary glands paired, acinous, although shown as 
tubular (probably erroneously) in Oenopota levidensis 
Shimek (1975). Venom gland of uniform histology. Muscular 
bulb with a thin outer muscular layer. 

Remarks. Species of this group were previously treated as 
Mangeliinae, but were isolated as a subfamily primarily on 
the basis of the presence of an operculum and a spirally 
sculptured protoconch (Bogdanov, 1987, 1990). None of 
these features are presently considered as being of subfamilial 
importance. However, one more character was revealed in 
our study, the structure of the salivary glands, which distin- 
guished Oenopotinae from the Mangeliinae. We provision- 
ally retain the subfamilial rank of Oenopotinae until the 
systematic importance of this character becomes certain. 



Subfamily Coninae Fleming, 1822 

Shell of medium to large size (usually 30-50 mm, up to more 
than 120 mm), biconic to conic. The inner shell walls are 
partially resorbed. Anterior canal short, aperture usually 
narrow, parallel-sided. Sculpture entirely spiral, usually weak 
or obsolete, sometimes tubercules on the shoulder. Anal 
sinus on the upper shoulder or almost sutural, shallow to 
relatively deep, occupying a rather narrow zone. Protoconch 
multispiral, smooth or spirally striated. Operculum small, 
with terminal nucleus, rarelyabsent. Egg capsules, bilaterally 
flattened, vasiform, arranged in clusters. 

Radula. Radular teeth harpoon-shaped, barbed or 
unbarbed on the tips, without solid base, usually open 
terminally (rarely laterally) at the base. 

FOREGUT. Proboscis moderately short and folded in con- 
tracted state. Rhynchostome lacks definite sphincter and 
rhynchodaeum can be greatly expanded to form a rostrum in 
fish-feeding species. Radial muscles lie in rhynchodeal wall. 



Subfamily Mangeliinae Fischer, 1884 

Shell small (usually 5-12 mm, up to 20 mm), ovate to 
fusiform. Anterior canal rather short. Both spiral and axial 
sculpture well developed. Anal sinus on the shoulder, shallow 
to rather deep, sometimes subtubular. Outer lip usually with 
terminal varix, sometimes denticulate. Protoconch smooth or 
variously sculptured. Operculum absent. Egg capsules dome- 
shaped, with an operculum. 

Radula. Radular teeth hollow with a solid base, sometimes 
with a semi-enrolled shaft; tooth canal opens laterally. 

Foregut. Proboscis moderately long, with a single or no 
distal sphincter, intermediate and posterior sphincters some- 
times present. Buccal tube introvert ('valvule') present. Dis- 
tal lips of buccal tube can be inverted. Buccal lips large and 
can be introverted into the buccal cavity. Buccal mass basal. 
Salivary glands paired and tubular, accessory salivary glands 
absent. Venom gland of uniform histology; muscular bulb 
usually with a thin outer muscle layer. 



162 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



Subfamily Daphnellinae Deshayes, 1863 

Small to moderately large shells (usually 5-15 mm, deep-sea 
species larger, up to 95 mm). Anal sinus sutural, shaped as a 
reversed-L, or on the upper shoulder and varying in depth. 
Sculpture variable, usually cancellate or with predominant 
spirals, and often with a smooth shoulder. Protoconch usually 
multispiral, rarely paucispiral, typically diagonally cancel- 
lated, although some genera have spiral or axial ribbing. 
Operculum absent. Egg capsules dome-shaped operculate. 

Radula. Radular teeth with large solid base and barbed or 
unbarbed tips, tooth cavity opens laterally at the base. 
Radula absent in some species. 

Foregut. Rhynchodeal introvert present in many species. 
Rhynchodeal septum present in some species. Proboscis 
usually short, often absent. Buccal mass basal. Radula appa- 
ratus absent in many species, vestigial in Gymnobela emer- 
toni. Radial muscles present in the rhynchodeal wall in 
radula- and proboscis-less species. Buccal lips well devel- 
oped, can be intverted into the buccal cavity. Salivary glands 
paired tubular or absent. Accessory salkivary glands absent. 
Venom apparatus absent in many species. In Daphnella 
reeveana the anterior part of venom gland is ciliated. Muscu- 
lar bulb can be single layered. 

Remarks. Although Thatcheria is sometimes classified in a 
separate subfamily Thatcherinae, we failed to find any ana- 
tomical or shell characters which would justify separation 
from the Daphnellinae. 



advised on problems of phylogenetic analysis. Yuri Kantor and John 
Taylor are grateful to the Royal Society, London and the Russian 
Academy of Sciences, Moscow for grants which enabled them work 
in London and Moscow respectively. Yuri Kantor and Alexander 
Sysoev gratefully acknowledge a grant from the Soros Foundation 
and the Russian Academy of Natural Sciences. 



APPENDIX 1 



Features of the shell 

Shell characters are still important for the systematics of 
Conoidea, and thus should be included in the analysis. 
However, there is probably no shell character which is 
diagnostic of any single group. Moreover, there has been no 
analysis of the adaptive or evolutionary significance of these 
shell features. Nevertheless, a few shell characters appear to 
be useful for the separation of clades. 

Shell shape 

This character which is concerned with overall shell shape is 
the most subjective. We recognise five basic shell shapes: 
1, fusiform shell; 2, cone-shaped shell; 3, turreted shell; 
4, terebriform shell; 5, a large group of 'intermediate' states, 
'biconic-fusiform', 'ovate-biconical', 'ovate-fusiform', 'clavi- 
form', etc. characterized by rounded outlines of the shell, 
which is more or less oval in its general profile. 



Subfamily Taraninae Casey, 1904 

Shell very small (up to 6 mm), ovate-fusiform. Anterior canal 
rather short. Sculpture well developed. Anal sinus very broad 
and shallow, situated on the shoulder or immediately below 
it. Protoconch paucispiral, finely spirally striated, or with 
spirally aligned granules. Operculum and radula absent. 

Foregut. Rhynchostomal sphincter absent, no radial 
muscles in rhynchodeal wall. Proboscis absent. Buccal mass 
undefined. Salivary glands absent. Venom apparatus absent. 

Remarks. This monotypic radula-less subfamily was rein- 
stated (Kantor & Sysoev, 1989) because it differs in shell 
characters from any other turrids lacking a radula. However, 
the very simplified morphology makes the evaluation of the 
status of the subfamily difficult. For the present we conserve 
the subfamily, but are unsure of its status. 



Acknowledgements. For gifts and loan of material without which 
this study could not have been completed, we are extremely grateful 
to J.H. McLean, the late V. Maes, R.N. Kilburn, F.E.Wells, I. 
Loch, R. L. Shimek, P. Bouchet, S. Gofas, B. Morton, W. F. 
Ponder, G. Rosenberg, and B. Marshall. Other material was 
obtained from the Institute of Oceanology, Academy of Sciences, 
Moscow; Zoological Institute, Academy of Sciences, St Petersburg, 
Zoological Museum, Moscow State University and we are very 
grateful to the curators L.I. Moscalev, A.N. Mironov, B.I Sirenko 
and D.L. Ivanov for their kind permission to use this material. David 
Cooper expertly prepared many of the serial sections of gastropod 
foreguts. Some unpublished material was made available by John 
Miller and A.I Medinskaya. The optical photomicrographs were 
made by Peter York. Andrew Smith and David Reid patiently 



Number of protoconch whorls 

Two types of protoconch can be recognised; the paucispiral 
and multispiral. These types of the protoconch were into- 
duced into turrid systematics by Powell (1942, 1966) and they 
are widely used in taxonomy. Generally, this subdivision 
coincides with that between planktotrophic and non- 
planktotrophic modes of larval development, although there 
are many exceptions to the rule among turrids (Bouchet, 
1990). The character is considered as being of little phyloge- 
netic importance (Bouchet, 1990), but a predominance of a 
single type of the protoconch can be noted in some taxa. For 
instance, most Daphnellinae and Conidae have multispiral 
protoconchs, whilst the paucispiral type is a typical of the 
Oenopotinae (Bogdanov, 1990). Protoconchs with 1-2 whorls 
are here considered as paucispiral, and these with two or 
more whorls as multispiral (Bouchet, 1990). 

Sculpture of the protoconch 

The pattern of protoconch sculpture has been widely used in 
conoidean taxonomy since Powell (1942, 1966). Turrids have 
a very wide variety of protoconch sculpture and at present, 
we are unable to classify them into clearly defined types. 
Thus we recognize only two major states of the character; 
firstly protoconchs lacking or with only weakly defined sculp- 
ture and secondly, protoconchs with well developed sculp- 
ture. Some higher taxa may be characterized by the presence 
or absence of protoconch sculpture. For example, the closely- 
related Turricula and Clavatula usually possess a smooth 
protoconch, whilst in the Turrinae it is usually axially costate. 
The only type of the protoconch sculpture characteristic of a 
single subfamily is the 'diagonally cancellated' form found 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



163 



among species of Daphnellinae, although this is not present 
in all species. 

Length of siphonal canal 

To define groups of shells with different lengths of the 
anterior siphonal canal we used parameter Rsl (relative 
siphonal length) of Harasewych (1981). As a result, we 
recognise three states of the character; a long siphonal canal 
(Rsl more than 0.39; up to 0.48 in the species studied), 
moderate canal (Rsl 0.21 to 0.34), short canal (Rsl less than 
0.20) or not differentiated from the apertural canal. 

Position of the anal sinus 

The anal (labial) sinus is a characteristic feature of Turridae 
and its position on the shell whorls is widely used for 
characterizing species and higher taxa (Powell, 1942, 1966; 
McLean, 1971). We follow Powell (1966) in recognizing 4 
types of sinus position; sutural (the deepest point of the sinus 
is situated near the suture), subsutural (on the whorl shoul- 
der), peripheral, and poorly pronounced (or very slight). 
Most turrids have a subsutural sinus; a peripheral sinus is 
characteristic for all Turrinae and some Clathurellinae 
(Bathytoma and related genera); a sutural sinus is common 
among the Daphnellinae. A weak, almost imperceptible sinus 
occurs occasionally in many subfamilies. It should be empha- 
sized that sinus types are recognized by growth lines, since 
the form of sinus at the outer lip of a mature shell may not be 
the same as that of the immature gastropod. 

Operculum 

The presence of an operculum is obviously the primitive state 
of the character. All 'lower' conoideans have a well devel- 
oped operculum. Among 'higher' conoideans, the operculum 
is absent in almost all Daphnellinae and Mangeliinae, but 
retained in the Oenopotinae and Conidae. In Clathurellinae 
(incorporating Borsoniinae), the operculum may be present, 
vestigial or absent, even in apparently closely-related genera 
(McLean, 1971). 



Position of opercular nucleus 

The opercular nucleus is usually situated in a terminal posi- 
tion at the tip of the operculum, but in the Clavatulinae and 
Turricula it is located medio-laterally. 



Presence of apertural armament 

The aperture of conoidean shells may be without ornament 
on the outer lip or columella, or they may bear weak to strong 
denticles, plications and folds. Armed apertures are found in 
the subfamilies Mangeliinae, Clathurellinae and Daphnelli- 
nae, and mostly amongst tropical shallow-water species. 



Number of teleoconch shell whorls 

We recognize three types of shells by this character. 1. shells 
with a small number of whorls (4 and less); 2. with an 
intermediate number of whorls (5 to 8); 3. with many whorls 
(9 and more) 



Presence of well developed subsutural ramp 

A subsutural ramp, (a morphologically distinct, often flat- 
tened part of the whorl profile immediately below the suture) 
may be either absent, or pronounced, in many subfamilies of 
Turridae. Usually, this character is clearly shown by a change 
in both spiral and axial sculpture in this region of the whorl. 



APPENDIX 2 

Genus-group taxa of recent Turridae S.L. 
(Compiled by A.V. Sysoev) 

The list presented below is of Recent taxa of the genus-group 
of Turridae s.l. distributed in respect to the classification 
adopted in the present paper. Since all the data concerning 
genera described before 1966 were given in Powell's (1966) 
monograph, type-species and bibliographic citations are 
included only for genera and subgenera described after 1966. 
Synonymy is also given only when it differs from that adopted 
by Powell. 

The classification used is to a great extent conservative; we 
avoid the description of new taxa and radical changes in the 
existing classification. As a result, some genera are of 
'unclear' taxonomic position and cannot be assigned, despite 
anatomical information, to any existing subfamily (Toxico- 
chlespira, for example). Some other genera (such as Genota) 
are only provisionally included into a certain subfamily. 

There are 337 valid Recent genera and subgenera. 

Family DRILLIIDAE Morrison, 1966. ICZN pending 

Agtadrillia Woodring, 1928 
Eumetadrillia Woodring, 1928 

Bellaspira Conrad, 1868 

Calliclava McLean, 1971 
Veliger 14(1): 117 
Cymatosyrinx palmeri Dall, 1919 
Cerodrillia Bartsch & Rehder, 1939 
Lissodrillia Bartsch & Rehder, 1939 
Viridrillia Bartsch, 1943 

Clavus Montfort, 1810 
Plagiostropha Melvill, 1927 
Cymatosyrinx Dall, 1889 
Drillia Gray, 1838 
ClathrodrUlia Dall, 1918 

Elaeocyma Dall, 1918 

Globidrillia Woodring, 1928 

Horaiclavus Oyama, 1954 
Anguloclavus Shuto, 1983 

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 9-10 

Mangilia multicostata Schepman, 1913 
Cytharoclavus Kuroda & Oyama in Kuroda, Habe & Oyama, 1971 

The sea shells of Sagami Bay: 213 

Pleurotoma (Mangilia) filicincta Smith, 1882 

Imaclava Bartsch, 1944 
Iredalea Oliver, 1915 
Kylix Dall, 1919 
Leptadrillia Woodring, 1928 



164 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



Neodrillia Bartsch, 1943 

Orrmaesia Kilburn, 1988 

Ann. Natal Mus. 29(1): 201-202 
Orrmaesia dorsicosta Kilburn, 1988 

Splendrillia Hedley, 1922 
Hauturua Powell, 1942 

Spirotropis G.O.Sars, 1878 

Syntomodrillia Woodring, 1928 

Tylotiella Habe, 1958 

lAcinodrillia Kilburn, 1988 
Ann. Natal Mus. 29(1): 223 

Acinodhllia viscum Kilburn, 1988 (s.d. Kilburn, 1988, 
Ann. Natal Mus. 29(2): 557) 

IDouglassia Bartsch, 1934 

IFenimorea Bartsch, 1934 

IParacuneus Laseron, 1954 

Family PSEUDOMELATOMIDAE Morrison, 1966 

Hormospira Berry, 1958 
Pseudomelatoma Dall, 1918 
(=Laevitectum Dall, 1919) 

Tiariturris Berry, 1958 

Family STRICTISPIRINAE McLean, 1971 

Cleospira McLean, 1971 

Veliger 14(1): 125 

Monilispira ochsneri Hertlein & Strong, 1949 
Strictispira McLean, 1971 

Veliger 14(1): 125 

Crassispira ericana Hertlein & Strong, 1951 

Family TURRIDAE H. & A.Adams, 1853 

Subfamily CLAVATULINAE Gray, 1853 

Benthoclionella Kilburn, 1974 
Ann. Natal Mus. 22(1): 214 
Benthoclionella jenneri Kilburn, 1974 

Clavatula Lamarck, 1801 

Clionella Gray, 1847 

Makiyamaia Kuroda in MacNeil, 1960 

Perrona Schumacher, 1817 

Pusionella Gray, 1847 

Scaevatula Gofas, 1989 

Arch. Molluskenk. 120(1/3): 16 

Scaevatula pelisserpentis Gofas, 1989 
Toxiclionella Powell, 1966 
Caliendrula Kilburn, 1985 

Ann. Natal Mus. 26(2): 442-443 

Latiaxis? elstoni Barnard, 1962 

Turricula Schumacher, 1817 

IMakiyamaia Kuroda in MacNeil, 1960 

Subfamily CRASSISPIRINAE Morrison, 1966 

Aoteadrillia Powell, 1942 

Austrodrillia Hedley, 1918 
Regidrillia Powell, 1942 



Belalora Powell, 1951 

Buchema Corea, 1934 

Calcatodrillia Kilburn, 1988 

Ann. Natal Mus. 29(1): 290-291 
Calcatodrillia chamaeleon Kilburn, 1988 

Carinodrillia Dall, 1919 

Carinapex Dall, 1924 

Ceritoturris Dall, 1924 

Conorbela Powell, 1951 

Conticosta Laseron, 1954 

Crassiclava McLean, 1971 
Veliger 14(1): 121 
Pleurotoma turricula Sowerby, 1834 

Crassispira Swainson, 1840 
Burchia Bartsch, 1944 
Crassispirella Bartsch & Rehder, 1939 
Dallspira Bartsch, 1950 
Gibbaspira McLean, 1971 

Veliger 14(1): 122 

Pleurotoma rudis Sowerby, 1834 
Glossispira McLean, 1971 

Veliger 14(1): 121 

Pleurotoma harfordiana Reeve, 1843 
Monilispira Bartsch & Rehder, 1939 
Striospira Bartsch, 1950 
(= Adanaclava Bartsch, 1950) 



Doxospira McLean, 1971 
Veliger 14(1): 124 
Doxospira hertleini Shasky, 



1971 



Epideira Hedley, 1918 
(=Epidirona Iredale, 



1931) 



Funa Kilburn, 1988 

Ann. Natal Mus. 29(1): 267-268 
Drillia laterculoides Barnard, 1958 

Haedropleura Bucquoy, Dautzenberg & Dollfus, 1883 

Hindsiclava Hertlein & Strong, 1955 
(= Turrigemma Berry, 1958) 

Inodrillia Bartsch, 1943 

Inquisitor Hedley, 1918 

Kurilohadalia Sysoev & Kantor, 1986 

Zoologicheskij Zhurnal 65(10): 1462-1463 
Kurilohadalia elongata Sysoev & Kantor, 1986 

Lioglyphostoma Woodring, 1928 

Maesiella McLean, 1971 
Veliger 14(1): 123 
Maesiella maesae McLean & Poorman, 1971 

Mauidrillia Powell, 1942 

Miraclathurella Woodring, 1928 

Naskia Sysoev & Ivanov, 1985 

Zoologicheskij zhurnal 64(2): 196-197 
Naskia axiplicata Sysoev & Ivanov, 1985 

Naudedrillia Kilburn, 1988 

Ann. Natal Mus. 29(1): 276-278 
Naudedrillia nealyoungi Kilburn, 1988 

Nquma Kilburn, 1988 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



165 



Ann. Natal Mus. 29(1): 247 
Pleurotoma rousi Sowerby, 1886 

Plicisyrinx Sysoev & Kantor, 1986 

Zoologicheskij Zhurnal 65(10): 1465-1466 
Plicisyrinx decapitata Sysoev & Kantor, 1986 

Psittacodrillia Kilburn, 1988 

Ann. Natal Mus.: 29(1): 253 
Pleurotoma bairstowi Sowerby, 1886 

Ptychobela Thiele, 1925 

Turridrupa Hedley, 1922 

IParadrillia Makiyama, 1940 

(= Iwaoa Kuroda, 1953) 

(= Vexitomina Powell, 1942) 

Coronacomitas Shuto, 1983 
Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 1-2 
Paradrillia (Coronacomitas) gemmata Shuto, 1983 

? Pseudexomilus Powell, 1944 

Subfamily ZONULISPIRINAE McLean. 1971 

Compsodrillia Woodring, 1928 
Mammillae drillia Kuroda & Oyama in Kuroda, Habe & Oyama, 
1971 

The sea shells of Sagami Bay: 208 
Compsodrillia { Mammillaedrillia) mammillata 
Kuroda & Oyama in Kuroda, Habe & Oyama, 1971 

Pilsbryspira Bartsch, 1950 
Nymphispira McLean, 1971 
Veliger 14(1): 126 
Crassispira nymphia Pilsbry & Lowe, 1932 

Zonulispira Bartsch, 1950 

Subfamily COCHLESPIRINAE Powell, 1942 

Abvssocomitas Sysoev & Kantor, 1986 

' Zoologicheskij Zhurnal 65(10): 1461-1462 
Abyssocomitas kurilokamchatica Sysoev & Kantor, 1986 

Aforia Dall, 1889 

Abyssaforia Sysoev & Kantor, 1987 

Veliger 30(2): 117 

Aforia (Abyssaforia) abyssalis Sysoev & Kantor, 1987 
Dallaforia Sysoev & Kantor, 1987 

Veliger 30(2): 115-116 

Irenosyrinx? crebristriata Dall, 1908 
Steiraxis Dall, 1895 

Anticomitas Powell, 1942 

Antimelatoma Powell, 1942 

Antiplanes Dall, 1902 

(= Rectiplanes Bartsch, 1944) 

Apiotoma Cossmann, 1889 

Carinoturris Bartsch, 1944 

Clavosurcula Schepman, 1913 

Cochlespira Conrad, 1865 

(=Ancistrosyrinx Dall, 1881) 

(=Pagodosyrinx Shuto, 1969 

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 19(1): 190-191 
Pleurotoma (Ancistrosyrinx) travancorica granulata Smith, 1904) 

Comitas Finlay, 1926 

Fusiturricula Woodring, 1928 
Fusisyrinx Bartsch, 1934 



Knefastia Dall, 1919 

Leucosyrinx Dall, 1889 
Sibogasyrinx Powell, 1969 
Indo-Pacific Moll. 2(10): 343 
Surcula pyramidalis Schepman, 1913 

Marshallena Allan, 1927 

Megasurcula Casey, 1904 

M/iowa MacNeil, 1960 

Paracomitas Powell, 1942 

Parasyrinx Finlay, 1924 
Lirasyrinx Powell, 1942 

Pyrgospira McLean, 1971 
Veliger 14(1): 119 
Pleurotoma obeliscus Reeve, 1843 

Rhodopetoma Bartsch, 1944 

Schepmania Shuto, 1970 Venus 29(2): 37-38 Surcula variabilis Schep- 
man, 1913 

'? Micropleurotoma Thiele, 1929 

Subfamily TURRINAE H. & A.Adams, 1853 (1840) 

Cryptogemma Dall, 1918 

Decollidrillia Habe & Ito, 1965 

Epidirella Iredale, 1931 

Fusiturris Thiele, 1929 

Gemmula Weinkauff, 1875 
Pinguigemmula MacNeil, 1960 
Ptychosyrinx Thiele. 1925 

Lophiotoma Casey, 1904 

(= Lophioturris Powell, 1964) 
Unedogemmula MacNeil, I960 
Xenuroturris Iredale, 1929 

Lucerapex Iredale, 1936 

Polystira Woodring, 1928 

Turris Roeding, 1798 
Annulaturris Powell. 1966 

Family CONIDAE Fleming, 1822 

Subfamily CLATHURELLINAE H. & A.Adams, 1858 

'bathytomid' group of genera 

Bathytoma Harris & Burrows, 1891 
Micantapex Iredale, 1936 
Parabathytoma Shuto, 1961 
Riuguhdrillia Oyama, 1951 

Paraborsonia Pilsbry, 1922 

'borsoniid' group of genera 

Asthenotoma Harris & Burrows, 1891 

Borsonella Dall, 1908 

Borsonellospis McLean, 1971 
The Veliger 14(1): 126-127 
Leucosyrinx erosina Dall, 1908 
Borsonia Bellardi, 1839 
Boettgeriola Wenz, 1943 

Cordieria Rouault, 1848 



166 

Cruziturricula Marks, 1951 

Ophiodermella Bartsch, 1944 

Tropidoturris Kilburn, 1986 

Ann. Natal Mus. 27(2): 645-646 
Pleurotoma scitecostata Sowerby, 1903 

Typhlomangelia G.O.Sars, 1878 

Typhlosyrinx Thiele, 1925 

IDarbya Bartsch, 1934 

'clathurellid' group of genera 

Clathurella Carpenter, 1857 

Comarmondia Monterosato, 1884 

Corinnaeturris Bouchet & Waren, 1980 
J. Moll. Stud.,suppl.8: 77 
Pleurotoma leucomata Dall, 1881 

Crockerella Hertlein & Strong, 1951 

Glyphostoma Gabb, 1872 
Glyphostomopsis Bartsch, 1934 
Euglyphostoma Woodring, 1970 
Prof. pap. U.S. Geol. Survey 306-D: 401 
Glyphostoma partefilosa Dall, 1919 

Nannodiella Dall, 1919 

Strombinoturris Hertlein & Strong, 1951 

lEtrema Hedley, 1918 
Etremopa Oyama, 1953 
Etremopsis Powell, 1942 

IGenota H. & A.Adams, 1853 

'mitromorphid' group of genera 

Anarithma Iredale, 1916 

Arielia Shasky, 1961 
Vexiariella Shuto, 1983 
Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 6 
Arielia (Vexiariella) cancellata Shuto, 1983 

Diptychophlia Berry, 1964 

Lovellona Iredale, 1917 

Maorimorpha Powell, 1939 

Mitrellatoma Powell, 1942 

Mitromorpha Carpenter, 1865 
Mitrolumna Bucquoy, Dautzenberg & Dollfus, 1883 
(=Apaturris Iredale, 1917) 
(=Cymakra Gardner, 1937) 
(= Helenella Casey, 1904) 
(= Ida Marwick, 1931) 
(= Mitrihara Hedley, 1922) 

Scrinium Hedley, 1922 

Zetekia Dall, 1918 

'tomopleurid' group of genera 

Drilliola Cossmann, 1903 

Microdrillia Casey, 1903 

(= Acropota Nordsieck, 1977, nom.nov. pro A crobela Thiele, 1925 
non Foerster, 1862 
The Turridae of the European seas: 59) 

Phenatoma Finlay, 1924 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 

Pulsarella Laseron, 1954 

Suavodrillia Dall, 1918 

Tomopleura Casey, 1904 
Maoritomella Powell, 1942 

lAustroturris Laseron, 1954 

Wilodrillia Hedley, 1922 

? Heteroturris Powell, 1967 

Indo-Pacific Moll. 1(7): 411 
Heteroturris sola Powell, 1967 

Subfamily? CONORBIINAE De Gregorio, 1890 

Conorbis Swainson, 1840 

Benthofascis Iredale, 1936 

Subfamily OENOPOTINAE Bogdanov, 1987 

Curdtoma Bartsch, 1941 
(= Widalli Bogdanov, 1986 
Zoologicheskij Zhurnal 65(1): 45 
Pleurotoma trevelliana Turton, 1834) 

Granotoma Bartsch, 1941 

Obesotoma Bartsch, 1941 

Oenopota Morch, 1852 

Nodotoma Bartsch, 1941 

Oenopotella Sysoev, 1988 

Zoologicheskij zhurnal 67(8): 1119-1120 
Oenopotella ultraabyssalis Sysoev, 1988 

Propebela Iredale, 1918 
Canetoma Bartsch, 1941 
(=Funitoma Bartsch, 1941) 

ILorabela Powell, 1951 

Subfamily MANGELIINAE Fischer, 1883 

Acmaturris Woodring, 1928 

Agathotoma Cossmann, 1899 

Anacithara Hedley, 1922 

Antiguraleus Powell, 1942 

Apispiralia Laseron, 1954 

Apitua Laseron, 1954 

Bactrocythara Woodring, 1928 

Bela Gray, 1847 

Belaturricula Powell, 1951 

Bellacythara McLean, 1971 
The Veliger 14(1): 128 
Clavatula bella Hinds, 1843 

Benthomangelia Thiele, 1925 

Brachycythara Woodring, 1928 

Cacodaphnella Pilsbry & Lowe, 1932 

Citharomangelia Kilburn, 1992 

Annals Natal Mus. 33(2): 508-9 
Mangilia africana Sowerby, 1903 

Clathromangelia Monterosato, 1884 

Cryoturris Woodring, 1928 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



167 



Cytharella Monterosato, 1875 

Cyrtocythara Nordsieck, 1977 
The Turridae of European seas: 34 
Pleurotoma albida Deshayes, 1834 

Rugocythara Nordsieck, 1977 
The Turridae of European seas: 35 
Pleurotoma rugulosa Philippi, 1844 

Eucithara Fischer, 1883 

Euclathurella Woodring, 1928 

Fehria van Aartsen, 1988 

La Conchiglia 20(232-233): 232 
Ginnania taprurcnsis Pallary, 1904 

Gingicithara Kilburn, 1992 

Annals Natal Mus. 33(2): 495-6 
Mangelia lyrica Reeve, 1846 

Glyphoturris Woodring, 1928 

Glyptaesopus Pilsbry & Olsson, 1941 

Guraleus Hedley, 1918 
Euguraleus Cotton, 1947 
Mitraguraleus Lascron, 1954 

Helerocithara Hedley, 1922 

Ithycyihara Woodring, 1928 

Kurtzia Bartsch, 1944 

Kurtziella Dall, 1918 
Granoturris Fargo, 1953 
Rubetlatoma Bartsch & Rehder, 1939 
Kurtzina Bartsch, 1944 

Leiocithara Hedley, 1922 

Lienardia Jousseaume, 1884 
Acrista Hedley, 1922 
Hemilienardia Boettger, 1895 
Thetidos Hedley, 1899 

Liracraea Odhner, 1924 

Macteola Hedley, 1918 

Mangelia Risso, 1826 

Mangiliella Bucquoy, Dautzenberg & Dollfus, 1883 
Lyromangelia Monterosato, 1917 

Marita Hedley, 1922 

Neoguraleus Powell, 1939 

Notocythurella Hertlein & Strong, 1955 

Papillocilhara Kilburn, 1993 

Annals Natal Mus. 33(2): 516-7 
Papillocilhara hebes Kilburn, 1992 

Paramontana Laseron, 1954 

Platycythara Woodring, 1928 

Pseudoetrema Oyama, 1953 

Pseudoraphitoma Boettger, 1895 

Pyrgocythara Woodring, 1928 

Saccharoturris Woodring, 1928 

Stellatoma Bartsch & Rehder, 1939 

Tenaturris Woodring, 1928 



Thelecythara Woodring, 1928 

Turrella Laseron, 1954 

Vitjazinella Sysoev, 1988 

Zoologicheskij zhurnal 67(8): 1122 
Vitjazinella multicostata Sysoev, 1988 

Vitricythara Fargo, 1953 

?.4rt»c7/m<raThiele, 1934 

IConopleura Hinds, 1844 

IHemicythara Kuroda & Oyama in Kuroda, Habe & Oyama, 1971 
The sea shells of Sagami Bay: 229 
Pleurotoma octangulaia Dunker, 1860 

1 Puraclathurella Boettger, 1895 

Subfamily DAPHNELLINAE Deshayes, 1863 

Abyssobela Kantor & Sysoev, 1986 
Zoologicheskij Zhurnal 65(4): 492 
Abyssobela atoxica Kantor & Sysoev, 1986 

Antimitra Iredale, 1917 

Asperdaphne Hedley, 1922 
Aspertilla Powell, 1944 

Austrodaphnella Laseron, 1954 

Bathybela Kobelt, 1905 
(= Bathypota Nordsieck, 1968 
The Turridae of European seas: 28 
Pleurotoma tcnellulu [sic) Locard, 1897) 

Buccinaria Kittl, 1887 

Cryptodaphne Powell, l l >42 
Acamptodaphne Shuto, 1971 
Venus 30(1): 10 
Pleurotomella biconica Schepman, 1913 

Cenodagreutes Smith. 1967 
The Vcliger 10(1): 1 
Cenodagreutes aetluts Smith, 1967 

Daphnella Hinds, 1844 
Diaugasma Melvill, 1917 
Hemidaphne Hedley, 1918 

Eubela Dall, 1889 

Eucyclotoma Boettger, 1895 

Exomilus Hedley, 1918 

Famelica Bouchet & Waren, 1980 
J. Moll. Stud.,suppl.8: 88 
Pleurotomella catharinae Verrill & Smith, 1884 

Fusidaphne Laseron, 1954 

Gymnobela Verrill, 1884 
(= Majox Nordsieck, 1968 

Die europaischen Meeres-Gehause Schnecken: 182 

Pleurotomella bairdi Verrill & Smith, 1884) 
(= Watsonaria Nordsieck, 1968 (nomen nudum) 

Die europaischen Meeres-Gehause Schnecken: 182 

Clathurella watsoni Dautzenberg, 1889) 
Theta Clarke, 1959 

lsodaphne Laseron, 1954 

Kermia Oliver, 1915 

Kuroshiodaphne Shuto, 1965 



168 



J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



Lusitanops Nordsieck, 1968 

Die europaischen Meeres-Gehause schnecken: 181 
Pleurotomella lusitanica Sykes, 1906 
(= Pseudazorita Nordsieck, 1977 (published as nomen nudum) 
The Turridae of the European seas: 31 (published as 
a subgenus of Thesbia) 

Pleurotoma blanchardi Dautzenberg & Fischer, 1896, 
s.d. Bouchet, Waren, 1980, 1980, J. Moll. Stud., 
suppl. 8: 83) 

Magnella Dittmer, 1960 

Microdaphne McLean, 1971 
The Veliger 14(1): 129-130 
Philbertia trichodes Dall, 1910 

Microgenia Laseron, 1954 

Neopleurotomoides Shuto, 1971 
Venus 30(1): 5-6 
Clathurella rufoapicata Schepman, 1913 

Nepotilla Hedley, 1918 

Ootomella Bartsch, 1933 

Pagodidaphne Shuto, 1983 

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 21 
Pagodidaphne colmani Shuto, 1983 

Philbertia Monterosato, 1884 
(= Lineotoma Nordsieck, 1977, nom.nov. pro Cirillia 

Monterosato, 1884 non Rondani, 1856 

The Turridae of the European seas: 18) 
Glyphostomoides Shuto, 1983 

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 16-17 

Philbertia (Glyphostomoides) queenslandica Shuto, 1983 

Phymorhynchus Dall, 1908 

Pleurotomella Verrill, 1873 

(= Azorilla Nordsieck, 1968 
Die europaischen Meeres-Gehause Schnecken: 184 
Pleurotoma megalembryon Dautzenberg & Fischer, 1896) 

(= Azorita Nordsieck, 1968 
Die europaischen Meeres-Gehause Schnecken: 184-185 
Pleurotoma bureaui Dautzenberg & Fischer, 1897) 

Anomalotomella Powell, 1966 

Ponliothauma Smith, 1895 

Pseudodaphnella Boettger, 1895 

Raphitoma Bellardi, 1848 
Cyrtoides Nordsieck, 1968 

Die europaischen Meeres-Gehause schnecken: 176 
Raphitoma rudis Scacchi, 1836 (= R. (C.) neapolitana Nords- 
ieck, 1977, nom.nov. pro R. rudis Scacchi, 1836 non Broderip) 

Rimosodaphnella Cossmann, 1915 

Spergo Dall, 1895 
Speoides Kuroda & Habe, 1961 

Stilla Finlay, 1926 

Tasmadaphne Laseron, 1954 

Teretia Norman, 1888 

Teretiopsis Kantor & Sysoev, 1989 
J. Moll. Stud. 55: 538 
Teretiopsis levicarinatus Kantor & Sysoev, 1989 

Thatcheria Angas, 1877 

Tritonoturris Dall, 1924 



Truncadaphne McLean, 1971 
The Veliger 14(1): 129 
'Philbertia' stonei Hertlein & Strong, 1939 

Tuskaroria Sysoev, 1988 

Zoologicheskij Zhurnal 67(7): 970-972 
Tuskaroria ultraabyssalis Sysoev, 1988 

Veprecula Melvill, 1917 

Vepridaphne Shuto, 1983 

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 17 
Daphnella cestrum Hedley, 1922 

Xanthodaphne Powell, 1942 

Zenepos Finlay, 1928 

lAliceia Dautzenberg & Fischer, 1897 

? Benthodaphne Oyama, 1962 

lOtitoma Jousseaume, 1898 

IThesbia Jeffreys, 1867 

Subfamily? TARANINAE Casey, 1904 

Taranis Jeffreys, 1870 

CONIDAE INCERTAE SEDIS 

Austrocarina Laseron, 1954 

Austropusilla Laseron, 1954 
Metaclathurella Shuto, 1983 

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 15 
Austropusilla (Metaclathurella) crockerensis Shuto, 1983 

Paraspirotropis Sysoev & Kantor, 1984 

Zoologicheskij Zhurnal 63(7): 1096-1097 
Pleurotomella simplicissima Dall, 1907 

Teleochilus Harris, 1897 

Toxicochlespira Sysoev & Kantor, 1990 
Apex 5(1-2): 2-3 
Toxicochlespira pagoda Sysoev & Kantor, 1990 

Typhlodaphne Powell, 1951 

CONOIDEA INCERTAE SEDIS 

Cretaspira Kuroda & Oyama in Kuroda, Habe & Oyama, 1971 
The sea shells of Sagami Bay: 219 

Cretaspira cretacea Kuroda & Oyama in Kuroda, Habe & 
Oyama. 1971 

Graciliclava Shuto, 1983 

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 11 
Graciliclava mackayensis Shuto, 1983 

Inkinga Kilburn, 1988 

Ann. Natal Mus. 29(1): 230 

Pleurotoma (Clionella) platy stoma Smith, 1877 

Kurodadrillia Azuma, 1975 

Venus 33(4): 159 

Kurodadrillia habui Azuma, 1975 
Lioglyphostomella Shuto, 1970 

Venus 28(4): 165-166 

Drillia timorensis Schepman, 1913 

Meggittia Ray, 1977 

Contribution to the knowledge of the molluscan fauna of 

Maungmagan, Lower Burma...: 66-67 
Meggittia maungmagana Ray, 1977 

Thatcheriasyrinx Powell, 1969 



FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



169 



Indo-Pacific Moll. 2(10): 405 

Ancistrosyrinx orientis Melvill, 1904 (by monotypy) 

Viridoturris Powell, 1964 (formerly Turrinae) 



Taxa transferred to other families 

Bathyclionella Kobelt, 1905 — Buccinidae (as synonym of 
Belomitra; Bouchet, Waren, 1980, J. Moll. Stud., suppl.8) 

Belomitra Fischer, 1882 — Buccinidae 

Steironepion Pilsbry & Lowe, 1932 — Columbellidae 

Surculina Dall, 1908 — Turbinellidae (Rehder, 1967, Pacific 
Sci. 21(2): 182-187) 

Turrijaumelia Sarasua, 1975 
Poeyana 140: 12-13 
Turrijaumelia jaumei Sarasua, 1975 

Transferred to Columbellidae as a synonym of Steironepion 
Pilsbry & Lowe, 1932 (Finlay, 1984, Nautilus 99(2-3): 73-75) 



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CO 



NTENTS 



97 The status of the Persian Gulf sea snake Hydrophis lapemoides (Gray, 1849) (Serpentes, 

Hydrophiidae) 

A. Redsted Rasmussen 
107 Taxonomic revision of some Recent agglutinated foraminifera from the Malay Archipelago, 

in the Millett Collection, The Natural History Museum, London 

P. Bronnimann and J.E. Whittaker 
125 Foregut anatomy, feeding mechanisms, relationships and classification of the Conoidea 

(= Toxoglossa) (Gastropoda) 

J.D. Taylor, Y.I. Kantor and A.V. Sysoev 



Must 



ZOOLOGY SERIES 

Vol 59, No. 2, November 1993 



ISSN 0968-0470 



Bulletin of 
The Natural Histor 
Museum 1 



THE NATURAL 
HISTORY MUSEUM 



Zoology Series 




THE 

NATURAL 
HISTORY 
MUSEUM 



VOLUME 60 NUMBER 1 23 JUNE 1994 



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© The Natural History Museum, 1994 



Zoology Series 
ISSN 0968 - 0470 Vol. 60, No. 1, pp. 1-104 

The Natural History Museum 

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London SW7 5BD Issued 23 June 1994 

Typeset by Ann Buchan (Typesetters), Middlesex 
Printed in Great Britain at The Alden Press, Oxford 



Bull. nat. Hist. Mus. Lond. (Zool.) 60(1): 1-37 



Issued 23 June 1994 



THE NATUR AL 
HISTORY MUSEU 

15 JUL 1994 
hicoENTtD 



A new subfamily and genus in Achatinidae 
(Pulmonata: Sigmurethra) 



ZOOLOGY LIB RA 



ALBERT R. MEAD 

Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA 

CONTENTS 



Synopsis 1 

Introduction 1 

Methods 3 

Abbreviations - Anatomical 3 

Achatinidae 3 

Key to subfamilies 3 

Callistoplepinae - new subfamily 3 

Key to genera 4 

Genus Callistoplepa 5 

Key to species 5 

Genus Leptocala 12 

Key to species 13 

Achatininae 18 

Genus Bequaertina - new genus 18 

Key to species 22 

Radulae and j aws 32 

Acknowledgements 35 

Acronyms 35 

Re f e rences 36 



Synopsis. In the Achatinidae, characteristics of the internal anatomy, particularly those in the reproductive tract, 
are continuing to show greater dependability in determining phylogenetic affinities than those in the shell, radula or 
jaw. In 1934, J.C. Bequaert reached the conclusion in the course of his revising the African land snail family 
Achatinidae, 'that several of the East African Achatinae are not separable from the West Africa Callistoplepae, at 
any rate as far as shell characters go'. The present 5-year study of over 500 shell specimens and 50 soft anatomies, 
involving 11 genus-group and 57 species-group nomina, demonstrates that the 4 anatomically distinct West African 
species, to which Bequaert referred, are the most primitive in the family. Accordingly, they have been placed in a 
separate subfamily. Anatomically the 5 contrastingly different East African species were found to be a distinct 
genus, plesiomorphic to subgenus Achatina [sensu Bequaert, 1950). Their phylogenetic relationships show a strong 
correlation with the distributional evidence in 60 recorded localities, which delineates (1) a short north central axis, 
(2) a central to eastern Africa axis, and (3) a strong north-south axis extending from eastern Africa into southern 
Africa. This evidence supports the emerging pattern of terrestrial gastropod distribution in Africa. The present 
project forms the foundation for a revision of the family, currently in progress. 



INTRODUCTION 



Fifty-seven species-group nomina associated with the acha- 
tinid genus Callistoplepa have provided the basis for the 
present 5-year project. The examination of over 500 shell 
specimens and 50 soft anatomies has revealed the fact that 
four West African anatomically distinct but related Lower 
Guinea species, in two genera, constitute a separate, primi- 
tive subfamily of the Achatinidae. Anatomical and concho- 
logical studies place the five other valid species in a new 
genus that is distributed in Central, Eastern and Southern 
Africa. 



Taxonomy in the Achatinidae is based both on the shell 
and on the soft anatomy, particularly that of the basal 
reproductive tract. The shell is a permanent record that 
reflects ontogenetically the influence of the environment, 
whereas the soft anatomy reflects phylogenetically the influ- 
ence of evolution. The latter, in the short term, is relatively 
free of substantive changes, being limited to temporary 
variations that reflect nutritional, developmental or repro- 
ductive influence. For example, both immature and seriously 
malnourished mature specimens have misleadingly attenu- 
ated, thin reproductive tracts with greatly limited muscula- 
ture; but the malnourished specimen can be readily 
distinguished because it has a much larger reproductive tract 



) The Natural History Museum, 1994 



A.R. MEAD 



with the rich coloration of the mature specimen. Under 
suitable conditions, both types of individuals are able to 
assume the basic genital proportions that are typical of the 
species. In the long term, the shell reflects and responds to 
the pressures of selection in a changing or in a new environ- 
ment. Wholly different molluscan stocks moving into distinct 
but similar environments may evolve convergently into forms 
that are so similar that conchologically they cannot reason- 
ably be distinguished or identified without first knowing the 
locality, e.g. Achatina (Lissachatina) craveni Smith, 1881 and 
A. (Achatina) tavaresiana Morelet, 1866 (Mead, 1992). 
Although in the long term, the soft anatomy undergoes 
changes, it revealingly does so within a more restricted, basic, 
generalized patten obviously characteristic of the larger 
group of which it is a related part. At this stage of investiga- 
tions, these larger groups seem to be taking shape within 
genera and subgenera (sensu Bequaert, 1950). 

All these factors have so entered into the present project 
that nine species, once thought to be congeners on the basis 
of the great similarity in their familial unique shell characters, 
are now separated into two subfamilies. The shells are 
remarkably similar, but the soft anatomies convincingly tell a 
different story. In essence, in the Achatinidae the soft 
anatomy reveals dependable, differentiating taxonomic crite- 
ria at the species level and above; the shell usually reveals 
supporting taxonomic criteria at the species level and often 
convincing criteria at the subspecies level. A more clear 
concept of subspecies is beginning to emerge in this family: 
Essentially consistent, usually minor shell differences in the 
members of an allopatric population that have features in 
their soft anatomy indistinguishable from those of the nomi- 
nate subspecies. This is precisely why the erstwhile enigmatic 
Leptocala petitia is shown in the text to be a distinct species 
rather than a subspecies or a synonym of the conchologically 
very slightly different L. mollicella. 

The shells of the nine species in this project have been 
redescribed in the light of larger series of specimens and more 
detailed examinations of shell characters than in previous 
studies. In many full grown specimens, weathering and 
environmental abrasion have obscured or removed valuable 
shell characters, particularly in the upper whorls. The empha- 
sis on a comparative examination of juvenile specimens in the 
present study, therefore, has been especially informative. In 
fact, it strongly supports the value of the collecting juvenile 
specimens, along with the larger specimens, whenever they 
are available. For the same reasons, determining the number 
of whorls is always an imprecise measurement and thus is 
recorded here only to the nearest one-quarter whorl. The 
length of the last whorl (= body whorl) includes the entire 
whorl and is measured from the base of the aperture to the 
point on the suture immediately above where the outer lip 
attaches at the periphery. All shell measurements are carried 
to the nearest 0.1 mm. For many decades the basic horizontal 
and vertical sculpture of the achatinid shell has been referred 
to as 'decussate'; this has been changed in the present work to 
the more accurate term 'cancellate'. From the shell dimen- 
sions, two important relationships are drawn: 1) between the 
greatest shell width and the shell length, and 2) between the 
length of the last whorl and the shell length. These relation- 
ships are expressed in percentages rather than in ratios. This 
emphasizes the measurement being compared rather than the 
measurement to which it is being compared; that is, in 
making comparisons, it is easier to comprehend that the last 
whorl is 87% of the shell length, than the fact that the length 



is 1.15 times the length of the last whorl. Aperture width does 
not include the callus. The descriptive shell terminology is 
largely based on Cox (1960). 

Diligent searching in 49 museums and personal collections, 
fortunately brought to light an unexpected fair number of 
alcohol preserved specimens. The museums with the largest 
and most varied collections of alcohol preserved achatinid 
specimens are in Tervuren, Stockholm, Berlin, Bruxelles, 
Paris, Frankfurt and London. But very valuable specimens 
have been found in museums where there is relatively limited 
material. It was through the convictions of Edmond Dart- 
evelle of the Museum National de l'Afrique Centrale (Ter- 
vuren) that the wet collection of the museum is unparalleled 
in quantity and diversity. Because the available material used 
in the present project is limited, determined efforts were 
made, after establishing the basic genital pattern, to conserve 
the remaining specimens for future investigators. 

The relationships of the basal genital system show to 
greater comparative advantage in ventral view. For this 
reason, the line drawings are similarly oriented in this per- 
spective, with the male conduit to the left and the female 
conduit to the right, unless otherwise indicated. The indi- 
vidual structures of the conduits are spread apart to show 
their configuration and minimize obstruction. The drawings 
are idealized, where possible through multiple specimens, to 
eliminate irrelevant and misleading features produced in 
preservation. The origin of the penial retractor is on the male 
conduit at a point that marks the division between penis and 
vas deferens. As the retractor anlage reaches apically during 
development, it usually inserts somewhere on the columellar 
muscle system, but may insert on the body wall, diaphragm, 
transverse myoseptum or other sites in the haemocoele 
(Mead, 1950). The configuration of the developing viscera 
may predispose the manner of insertion. The not infrequent 
bifurcate (Fig. 20) and multifurcate penial retractor insertions 
support the ontogenetic rather than the functional interpreta- 
tion of these terms. Within a species, the site of insertion may 
be consistent or variable. There is a fairly strong tendency in 
this family for the basal penial retractor to proliferate muscle 
and apparently connective tissue that variously produce adhe- 
sions in the several parts of the male conduit. This in turn 
changes the relationship of these parts, alters the extrover- 
sion process, and the configuration of the resultant intromit- 
tent organ. This organ is normally composed seriatim, apex 
to base, of the penis, pilaster (when present), penis sheath, 
penial atrium and genital atrium externally, and the basal vas 
deferens, penial retractor, apical vas deferens and ejaculatory 
duct internally. 

Directions of left and right refer to those of the snail. 
Apical, in reference to the genital system, means toward the 
ovotestis, basal toward the genital atrium. The anatomical 
terminology is essentially that of Mead (1950) (see Abbrevia- 
tions - Anatomical below). 

The discussion of each species is in the following format: 
shell, soft anatomy (where available), type material, type 
locality, distribution and, when applicable, remarks. The 
sources and localities of the alcohol-preserved specimens are 
reported in the text. A table for each species includes the 
sources of all shell specimens examined, taxonomically 
important specimens, illustrated specimens, a size range, 
their localities, shell dimensions and shell proportions. Illus- 
trations in the literature are cited in the synonymies; the 
nature of the illustration, where it is other than of the shell, is 
shown in parentheses. Most localities were found in the 






NEW SUBFAMILY AND GENUS ACHATINIDAE 



volumes of the U.S. Board of Geographic Names and their 
locality figures were preferentially used. A list of acronyms of 
institutions and private collections follows the text. Symbols 
and abbreviations used in the Tables: Holo = Holotype, L = 
Length, Lect = Lectotype, LW = Last Whorl, Para = 
Paratype, PLec = Paralectotype, W = greatest Width, f = 
dissected, * = see photograph. 

Continued research along the lines of Mead (1950, 1978, 
1992) and in the present work promises to establish a sound 
taxonomic and phylogenetic base for the Achatinidae. 



METHODS 

Examining a great number of variously preserved whole 
specimens in the present project has once again emphasized 
the importance of using proper preservation procedures. 
Ideally, specimens selected for preservation of the soft parts 
should be put in previously boiled water that has reached 
ambient temperature. They should drown normally 8-12 
hours. Overdrowning will cause the basal genital structures to 
evert, irreparably distorting the taxonomically valuable fea- 
tures. Underdrowning permits the specimen to withdraw 
excessively into the shell. Crowding the specimens or using 
too small a volume of fluid promotes maceration. The 
adequately drowned specimen usually contracts slightly when 
placed in the initial 40% alcohol. In a few hours, depending 
upon the size of the specimen, it should next be placed in a 
60% solution, followed by at least one change to 70% 
alcohol. 

Formalin is a powerful, penetrating, irritating fixative. If it 
is used at a very low percentage for a short period before the 
specimen is washed thoroughly and transferred to 70% 
alcohol, it can be quite effective. But in general, its use 
should be avoided because formalin alters the colour of the 
shell, makes the shell brittle and chalky, causes the perios- 
tracum to crack and peel off upon drying, promotes adhe- 
sions between the shell and soft parts, severely hardens the 
muscular body wall, and precipitates great quantities of 
albumin and recrystallized calcareous islands in the tissues. 
As a result, extrication rarely can be accomplished without 
damage to both shell and soft parts. Further, even with 
prolonged soaking in a 0.5% trisodium phosphate solution, 
the muscular body wall remains so hard and tough that, with 
extreme difficulty it has to be snipped out, piece by small 
piece, to get to the soft parts, which often are so intensely 
fixed that they are brittle. Once the soft parts are removed, 
they usually can be relaxed for limited manipulation only by 
further soaking in trisodium phosphate and one or more 
prolonged water baths. Even then, a distressing degree of 
brittleness remains. The practice of 'neutralizing' formalin for 
the preservation of vertebrate specimens by first dropping 
live snail specimens in the solution is deplorable. 



ABBREVIATIONS - ANATOMICAL 



GA 


genital atrium 


OTD 


ovotestis duct 


P 


penis 


PA 


penial atrium 


PIL 


pilaster 


PR 


penial retractor 


PS 


penis sheath 


RCR 


right columellar retractor 


ROR 


right ommatophore retractor 


S 


spermatheca 


SD 


spermathecal duct 


SO 


spermoviduct 


ssv 


secondary seminal vesicle 


T 


talon 


V 


vagina 


VA 


vaginal atrium 


VR 


vaginal retentor 


Achatinidae 



AVD 


apical vas deferens 


BVD 


basal vas deferens 


E 


egg 


EM 


eversion muscle bands 


FO 


free oviduct 



Basal genital conduits simple, without accessory organs. A 
conspicuous sheath partially, or usually, completely envelops 
the penis. Spermatophores not formed. The right branch of 
the columella muscle system regularly remains to the left of 
the genitalia. Kidney long, two to three times the length of 
the pericardium; sigmurethrous. Pulmonary vein without 
major branches. Holopod. Rachidian tooth very slender and 
apparently nonfunctional, rarely wide. Jaw simple; smooth or 
usually striated. Shell ovate, elongate-ovate or conic-oblong, 
rarely columnar; anomphalous or umbilicate; columella trun- 
cate or continuous with outer lip, some forms are intermedi- 
ate. Endemic in continental Africa and its adjacent small 
coastal islands; four known introduced species elsewhere in 
the world. 

Key to Subfamilies 

Vas deferens does not penetrate the penis sheath, but leaves 
apically with the penial retractor through the sheath aperture. 
The penial retractor inserts on the right columellar retractor; 
it is extremely short, entirely or almost entirely covered by 
the penis sheath; penis contains a large, conspicuous pilaster. 

Rachidian tooth about as wide as the laterals CALLIS- 

TOPLEPINAE 

Vas deferens penetrates the penis sheath. Even within a 
single population, the penial retractor may variously insert on 
muscle bands, body wall, diaphragm or fascia; it is usually 
long to very long and entirely or almost entirely free of the 
penis sheath; penis contains an ill-defined pilaster, no pilas- 
ter, or a verge. Rachidian tooth usually much narrower than 
the laterals ACHATININAE 



CALLISTOPLEPINAE - new subfamily 

This subfamily contains the most primitive achatinids yet 
known. Phylogenetically, it is at the base of the Achatinidae, 
near the Subulinidae. Like that in the Subulinidae, the vas 
deferens does not penetrate the penial sheath. The more 
ovate, patterned shell, however, with its larger aperture and 
limited number of whorls places this taxon in the Achatinidae 
rather than the Subulinidae. Supportive of this are the long 
kidney and the pattern of lung venation. The radula is 
uncharacteristic of either family, but this is of lesser impor- 



A.R. MEAD 



tance phylogenetically because this structure is well known to 
be responsive to changing feeding habits within closely 
related species. The wide, functional rachidian tooth immedi- 
ately distinguishes the Callistoplepinae from most Achatini- 
nae (Fig. 58-63). D'Ailly (1896:69) was the first to examine 
and illustrate the radulae of both species of Callistoplepa (see 
also Pilsbry, 1904:ix,xv; Thiele, 1929:560 and Ortiz & Ortiz, 
1959:46). In the present study, the radulae of C. barriana 
(Sowerby, 1890), C. shuttleworthi (Pfeiffer, 1856) and Lepto- 
cala mollicella (Morelet, 1860) were found to follow the same 
basic form and pattern. Similarly all the jaws are essentially 
identical - simple, nearly smooth, fulvous, chitinoid collari- 
form band that is somewhat wider in the middle and tapering 
at the sides. There is no suggestion of even generic difference 
in these structures. Thus, with only two alcohol specimens of 
L. petitia (Jousseaume, 1884) extant, a decision was made to 
leave their odontophores intact. 

All four species in the Callistoplepinae are limited geo- 
graphically (ca 5° N-5° S) to the tropical Lower Guinea 
region of West Africa, which at this point appears to be the 
cradle of the Achatinidae (Mead, 1992). The high natural 
luster of their translucent, elongate-ovate shells probably 
reflects selective advantage in the protractedly wet rainforest. 
Prior to the present study, the soft anatomy had been 
examined in one or more species in the eleven achatinine 
genera except the Guinean genus Columna (none so far 
available) and all except four of the thirteen achatinine 
subgenera (sensu Bequaert, 1950). All that have been exam- 
ined clearly are anatomically more advanced than the two 
callistoplepine genera. 

The calcareous, thick-shelled eggs are comparatively large 
for the family and are on a par with those of Tholachatina. 
D'Ailly (1896:68) felt they were somewhat small compared to 
the size of the snail shell. Thiele (1929:560) echoed this point; 
but d'Ailly apparently was comparing them with the rela- 
tively huge eggs of some Archachatina and the subulinids. A 
reticulate-microtuberculate texture covers the slender, 
attenuated body. Basally, the mantle is generously covered 
with variable size, fusing black to gray spots; these show 
through the thin shell. Apically, the spots are smaller, more 
regular and concentrated on the shoulder of the whorls. The 
genital orifice appears to be unusually far posterior; Ortiz & 
Ortiz (1959) made this observation in C. shuttleworthi. 

Typical of the known achatinids, the anterior aorta in the 
Callistoplepinae is found on the dorsal surface of the dia- 
phragm where it abruptly penetrates the diaphragm to pass 
vertically along the anterior edge of the sagittal myoseptum. 
Anteriorly, this latter separates the right and left columellar 
retractors and incompletely places the male and female basal 
conduits in left and right chambers, respectively. In all four 
species, the triangular kidney is long, 2-3 times the length of 
the pericardium, and sigmurethrous. The ascending limb of 
the urethra is closed for its entire length. Venation of the lung 
is a dense, broad network on the pericardial side of the 
slender principal vein, whereas on the right of the principal 
vein there is a relatively narrow band of parallel, limitedly 
branching veins between it and the ascending limb of the 
ureter. The second largest vein, about half the caliber of the 
principal vein, starts as a network in the far left posterior 
corner of the lung, anastomoses, and joins the principal vein 
at a right angle 5-7 mm anterior to the kidney. 



Together, the four included species in this subfamily mani- 
fest seriatim, from simple to complex, an impressive transi- 
tion series in the basal male conduit from what surely is a 
pilaster to that which appears to be a verge or penis papilla. 
Callistoplepa barriana is the most primitive with an elongate, 
elevated pilaster on the ventral penial wall. In C. shuttlewor- 
thi the apical penis has permanently partially evaginated, 
pulling dorsally the basal-most part of the vas deferens into 
the pilaster and fixing it in place with tissue derived from the 
adjacent penial retractor. This progression is taken one step 
further in Leptocala petitia, wherein the permanently evagi- 
nated apical penis becomes so greatly enlarged, thick-walled 
and dorsoventrally distorted that the resultant pilaster essen- 
tially fills the thin-walled, saccular basal penis. Finally in L. 
mollicella, the pilaster assumes an apical position wherein it is 
axially pendulous within the thick-walled basal penis. At first 
glance, it appears to be a penis papilla, but the asymmetry 
within betrays the fact that it is in actuality a greatly modified 
pilaster. In all four species, the penial retractor is extremely 
short and inserts on the right columellar retractor. In C. 
shuttleworthi it inserts forward near the other branches; in the 
other species it inserts far to the rear. The penis sheath is so 
thin that it is difficult to trace; but it naturally enshrouds the 
entire penis, allowing the penial retractor and the vas defer- 
ens to pass out apically through the aperture of the sheath. A 
barely discernible transparent tissue layer attached directly 
on the surface of the penis is formed by the penial retractor. 
This may be the forerunner of the condition found in some 
Angolan achatinids, e.g. Achatina welwitschi Morelet, 1866, 
in which the penis is buried in dense muscle tissue extending 
from the penial retractor. Both the inner surface of the penis 
sheath and the adjacent but continuous outer surface of the 
penis are smooth, shiny and free from each other. This 
condition facilitates seriatim extroversion. 

The spermatheca is consistently attached to the spermovi- 
duct well above the junction of the apical vas deferens and 
the free oviduct. In the adult forms, there is no distinct 
vaginal retentor between the vagina and the body wall. In 
juvenile specimens of C. shuttleworthi, however, the anlage is 
present, which suggests that in this subfamily, its full develop- 
ment may be obviated by the highly developed, muscular 
basal female conduit. 



Type genus: Callistoplepa Ancey, 1888. 



Key to Genera 

Sculpture of body whorl coarse, with slender, tightly and 
evenly placed prosocline costate ridges; shell aperture large, 
usually 52% of shell length; last whorl long, usually > 73% of 
shell length. Posterior foot with dorsolateral serrate ridges. 
No colored band on neck. Equatorial Guinea to Nigeria- 

Callistoplepa 

Sculpture of body whorl with extremely finely engraved 
microscopic rhomboids or vertical vermiculate granulae; shell 
aperture modest, usually > 52% of shell length; last whorl 
shorter, usually > 73% of shell length. Foot without dorsolat- 
eral ridges. Dark gray band on neck between ommatophores 
and mantle. Cameroon to western Zaire Leptocala 






NEW SUBFAMILY AND GENUS ACHATINIDAE 



Callistoplepa 

Callistoplepa 

Ancey, 1888:69 (footnote 2 for 'Achatina shuttleworthiana' 
[sic = A. shuttleworthi Pfeiffer, 1856]); Pilsbry, 1905:viii, 
ix (fig. 2), xv (radula); Germain, 1909:90; Pilsbry, 
1919:54, 60, 80, fig. 25 (map); Bequaert & Clench, 
1934c:114; Ortiz & Ortiz, 1959:44; Zilch, 1959:372; Mead, 
1986:144. 

Ganomidos 

d'Ailly, 1896:66. Type species by present designation, 
Achatina barriana Sowerby, 1890. 

Callistopepla 

Ancey, 1898:92 (type species: Achatina shuttleworthi Pfe- 
iffer, 1856); Thiele, 1929:560; Germain, 1936:151 (foot- 
note 3); Verdcourt, 1966:111; Meredith, 1983:30; Oliver, 
1983:9; Parkinson, etal 1987:68; Vaught, 1988:90. 

Ganomidus 

Boettger, 1905:170. 

Ganomides 

Verdcourt, 1966:111. 

Callistoplepa s.s. 
Mead, 1992. 

After an extended trip to West Africa, a Captain Vignon 
prepared a catalogue of 104 land and freshwater molluscs that 
he had collected. The shells and the catalogue were subse- 
quently acquired by a collector in Marseille and made avail- 
able to his colleague C.F. Ancey, who was given the 
opportunity to publish this catalogue. Ancey (1888) agreed to 
present it 'such as it is, but with necessary, even indispens- 
able, annotations because of defective identifications, some 
of which are not found to be at the level of the science' 
(trans.). In one of many footnote annotations, he placed 
"Achatina shuttleworthiana' [sic] under a new generic name 
'Callistoplepa'. If Ancey was not responsible for the misspell- 
ing of the specific name 'shuttleworthi\ then at least he did 
not correct it. The greater misfortune was that he misspelled 
the proposed generic name, which as revealed later (Ancey, 
1898) was intended to be 'Callistopepla' (Gr. most beautiful 
robe). 

In view of Ancey's casual manner of publishing the descrip- 
tion of this genus, the spelling of the generic name 'Callis- 
toplepa" must be considered to be the 'correct original 
spelling' (ICZN Art. 32 (b) and is 'to be preserved unaltered'. 
According to Art. 32 (c), Ancey's name does not qualify as an 
'incorrect original spelling' because, 'without recourse to . . . 
external source of information,' there is no 'clear evidence of 
an inadvertent error' even though orthographically it would 
have been desirable to have spelled it 'Callistopepla' . His 
unorthodoxy and failure to make a timely correction in 
spelling suggested that he was content for ten years to leave it 
in its original form. In the meantime d'Ailly (1896) unwit- 
tingly proposed the generic synonym Ganomidos including 
Achatina shuttleworthi along with A. barriana. Further, 
Ancey's original spelling contravenes no provisions of the 
Code articles. It is only in his belated publication (1898) that 
he used the spelling 'Callistopepla' , without even implied 
justification for the change in spelling. Under the circum- 
stances, this constituted an 'unjustified emendation' of the 
original spelling and therefore it is a junior objective syn- 
onym (Art. 33 (b)(iii). Or, perhaps it was just another one of 
his regrettable misspellings. This rationale supports Pilsbry's 
conclusions (1905:126), but not those of Germain (1936:151 



footnote 3 ). Unfortunately, the confusion about the valid 
spelling of the generic name has persisted in collections and 
even in the more recent literature, e.g. Parkinson et ai, 
1987:68, Vaught, 1988:90. It is hoped that the present expli- 
cation finally will obviate any further confusion. 

Ancey (1888) gave as the outstanding characteristics of this 
new genus its totally different appearance, thin shell, fine 
striation, and a colour pattern recalling Orthalicus gallinasul- 
tana. d'Ailly (1896) was the first to describe adequately this 
taxon, emphasizing the delicate, translucent, shiny, white- 
flecked shell, the vertical filiform sculpture, the mammillate 
apex, the inflated body whorl, the elongate, serrate-cristate 
foot, the hard-shelled eggs, and the unusual radulae of both 
Ganomidos shuttleworthi and the then, newly embraced G. 
barriana. Pilsbry (1905) accepted broadly d'Ailly's character- 
ization of the genus and emphasized the importance of the 
very thin shell, the closely 'ribplicate' sculpture and the broad 
central tooth of the radula. In addition, he included in 
Ancey's genus Callistoplepa: Ganomidos pellucidus Putzeys, 
1898, G. fraterculus Dupius & Putzeys, 1900, Achatina mar- 
teli and its subspecies A. m. pallescens Dautzenberg, 1901. 
Germain (1909) and Pilsbry (1919) retained this grouping. 
Bequaert & Clench (1934c) added to this genus on the basis 
of shell characters: Achatina nyikaensis Pilsbry, 1909 and A. 
graueri Thiele, 1911. In the present work it is demonstrated 
on the basis of the soft anatomy that the taxa added to 
Callistoplepa since d'Ailly (1896) are not congeneric, but are 
in subfamily Achatininae. 

Bequaert & Clench (1934c: 114) were misleading when they 
reported that C. barriana and C. shuttleworthi 'are from 
Upper Guinea'. Columbia Lippincott Gazetteer (1952) 
defines Guinea as equatorial West Africa from Senegal to 
Angola, being divided into Upper and Lower Guinea by the 
Niger Delta, van Bruggen (1989) supports the interpretation 
that the division is at the Dahomey Gap. In either interpreta- 
tion, these species are limited to Lower Guinea. d'Ailly 
(1896:70) states that both species live in small numbers in 
shady places at the base of tree trunks and under detached 
pieces of bark. 

Key to Species 

Second whorl with thin crescentic threads and granules; last 
whorl evenly convex, expanding greatly, four times the length 
of the penultimate whorl when viewed dorsally; aperture 
length > shell width except in smallest specimens; peripheral 
arrow-shaped pattern usually pale and diffuse, occasionally 
absent or nearly so; the suture transects a smaller and often 
darker pattern; white flecks sparse or abundant, irregularly 
distributed; nepionic whorls 3; larger species (6 whorls = 
38-52 mm long). Genital aperture complex, large, superfi- 
cial; penial retractor inserts on the right columellar retractor 
posterior to all other branches; penis tubular; vagina longer 

than wide. Cameroon, Nigeria barriana 

Second whorl grossly deeply closely and evenly costate; last 
whorl subcarinate, expanding proportionately, three times 
the length of the penultimate whorl when viewed dorsally; 
aperture length < shell width; conspicuous light castaneous 
arrow-shaped pattern at periphery, with concentrations of 
white flecks tending to alternate with the pattern; a smaller 
similar pattern appears subsuturally, but the white flecks 
there are more scattered; nepionic whorls 2V2; smaller species 
(6 whorls = 26-34 mm long). Genital aperture simple, small, 
lacunate; penial retractor inserts on the right columellar 



A.R. MEAD 





BVD 




PIL 



RCR 








Fig. 1 Callistoplepa barriana, basal genital structures (MRAC no. 795.956). 

Fig. 2 C. barriana, right branch of columellar muscle showing posterior attachment of the penial retractor. 

Fig. 3 C. barriana, right ventrolateral view of penis to show the pilaster in profile. Contraction during preservation has telescoped the apical 

penis and forced it and its fibromuscular matrix out of the penis sheath. 
Fig. 4 Callistoplepa shuttleworthi, penis sheath, permanently partially evaginated penis, and pilaster (containing the basal vas deferens) are 

shown in frontal plane and in dorsal view (UUZM). 
Fig. 5 Same, in a slightly tangential sagittal plane. 

Fig. 6 Same, in ventral view with penis sheath cut longitudinally and spread laterally to show the penis within. 
Fig. 7 Same, with penis cut longitudinally and spread laterally to expose the pilaster within. Penis sheath not shown. 
Fig. 8 Same, with pilaster cut and spread to reveal the basal vas deferens opening dorsally into the lumen of the penis. The dense 

fibromuscular webbing at the junction of the basal vas deferens and penis has been removed for clarity. 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



retractor anterior to the retractor of the right optic tentacle; 
penis permanently partially evaginated; vagina wider than 
long. Cameroon, Gabon, Equatorial Guinea ('Grand Bas- 
sam' locality is suspect) shuttleworthi 

Callistoplepa barriana (Sowerby, 1890) 

Figs. 23, 24 

Achatina barriana 

Sowerby, 1890:579, pi. 56, fig. 2; von Martens, 1891:30. 
Ganomidos barrianum 

d'Ailly, 1896:70, pi. Ill, figs. 5-10 (egg), text fig. (radula). 
Callistoplepa barriana 

Pilsbry, 1904-05:127, pi. 47, figs. 14-17 (egg), pg. ix fig. 2, 

pg. xv (radula, ex d'Ailly); Germain, 1909:90; Bequaert & 

Clench, 1934c: 114. 
Ganomidus barrianum 

Boettger, 1905:170. 
Callistopepla barriana 

Dautzenberg, 1921:98; Oliver, 1983:9 (syntype). 

Shell. Shell ovate-conic, very thin, fragile, translucent, 
shiny. Whorls 6-6V4, rarely 6V2, moderately convex. The 
second and third nepionic whorls are nearly straight sided, 
but they immediately give way to postemergent rapidly 
expanding whorls, producing a mammillate or submammill- 
ate, broadly conic spire and a blunt apex. Shallow sutures 
form a thin, nearly even line. Last whorl large, convex, 82% 
of shell length, range for 4-6V2 whorls = 78-86% (n = 115), 
swelling faintly outward directly below the suture in some 
specimens. Aperture broadly ovate, nearly vertical, pale 
milky within. Columella thin, slender, slightly to broadly 
arcuate, concolorous, squarely to obliquely truncate, inner 
rim rolled adaxially. Outer lip thin, nearly evenly arcuate; 
joining the periphery at only a modestly acute angle; greatest 
width is characteristically midway. Parietal callus scarcely 
apparent in unweathered specimens. 

From apex to base, the shell ground collar is uniformly pale 
fulvous. Superimposed on this, beginning imperceptibly in 
the fourth whorl, are two narrow bands of slender yellow- 
brown chevrons - one at the periphery, and a less distinct one 
transected by the suture. The chevrons in close juxtaposition 
have their apices oriented prosocline and are about as wide as 
the space between them. Much thinner, more irregular, 
paler, parallel sinuate stripes may join the two bands. Speci- 
mens with the most conspicuous patterns may have a second 
zone of thin, pale, transverse bands between the periphery 
and the base of the shell. The peripheral pattern tends to fade 
with increased growth. Some specimens may have present 
only the sutural band, or a unicolorous last whorl, or an 
entirely unicolorous shell except possibly for a slightly darker 
transverse band laid down between growth periods. Any of 
the whorls may be flecked with minute circular or elongate 
white spots (usually ca 0.2-0.8 mm). These are irregularly 
and sparsely dispersed, but are especially conspicuous within 
the costae of the last whorl. Upon close examination, they are 
seen to be a consolidated white powdery substance between 
the two periostracal layers. Although some are associated 
with shell injuries, their formation is apparently a natural 
phenomenon contributing to cryptic coloration. 

The most apical portion of each nepionic whorl dips 
abruptly at near-right angles adaxially to form a narrow 
platform in which is embedded a strikingly uniform series of 
minute shallow pits that fringe the suture. This ornamenta- 



tion is limited to the nepionic whorls and is the homologue of 
the diagnostic grossly costate sculpture in the second nepionic 
whorl of C. shuttleworthi. The first whorl is essentially 
smooth. Short faint slender crescentic threads and granules, 
oriented transversely but aligned spirally in irregular series, 
gradually make their appearance in the second whorl. As this 
sculpture becomes more organized, the spaces between the 
several spiral series seem to form shallow spiral striae. Near 
the junction of the third and fourth whorls, a sharp transverse 
delineation marks the end of the nepionic whorls, at which 
level the threads become more symmetrical and greatly 
compressed, but retain their individuality. With continued 
growth, the threads remain fairly distinct or become trans- 
versely variously fused into costellae, which interrupt or 
obliterate in part the shallow spiral striae. Gradually the 
threads become more bold and evolve into slender, closely 
and very evenly placed prosocline corrugations or costae, 
commonly with splitting and anastomosis. The spiral striae 
remain superficial, barely transecting the costae. The 
depressed cancellate sculpture below the periphery of the 
upper whorls gradually becomes more corrugate until an 
essentially uniformly costate sculture is finally formed on the 
entire forward last whorl of the fullgrown specimen, dimin- 
ishing slightly toward the columella and obliterating the 
peripheral line of demarkation. The smallest shells may be 
vaguely subcarinate. 

Soft anatomy. Alcohol preserved specimens available 
31/dissected 13. Nigeria: BMNH 1/1; Cameroon: MRAC 2/2, 
SMNH 10/4, SMF 14/6, UUZM 4/0. d'Ailly (1896) had access 
to 34 alcohol-preserved specimens collected in Cameroon by 
P. Dusen, Y. Sjostedt and J.R. Jungner. With the generous 
assistance of Dr Ake Franzen, a diligent search was made in 
the museums of Stockholm and Uppsala in 1987, but only 14 
specimens could be found. There was no evidence of Jungn- 
er's specimens. 

The body of the preserved specimen is uniformly grey 
fulvous, without any apparent markings. Immediately poste- 
rior to the shell, there is a depressed plateau that is fringed by 
two prominent dorsolateral ridges, each composed of 12 
closely aligned, truncate incisor-shaped elevations. 

The most unusual feature of the internal anatomy of this 
species is the penis sheath (PS) (Fig. 1). Thickest at its base 
(~0.5mm) it diminshes apically to a diaphanous facia 
(~0.05 mm) that, in the normal position, enshrouds the 
apical penis (P), the most basal part of the vas deferens 
(BVD), and the basal portion of the extraordinarily short 
penial retractor (PR). As in other achatinids, the origin of the 
PR marks the division between P and BVD. In contrast to 
that in C. shuttleworthi, the PR inserts on the right columellar 
retractor (RCR) posterior to all other branches (Fig. 2). In 
the fully mature specimen (Fig. 1) the tapering attenuated 
apical P appears to be cramped into a sigmoid fold in this 
thinnest apical PS. A dense webbing of muscle and connec- 
tive tissue fibrils, originating from the PR, obscures, 
entangles and foreshortens the apical folds of the P, even to 
the point in the oldest specimens where this tight, wooly mass 
of fibromuscular tissue becomes histologically intimately 
intermeshed with the substance of the apical penial wall. On 
its outer surface, this cocoon-like network forms a smooth, 
dense coating over the P that is completely free from the 
equally smooth but very shiny inner surface of the PS, thus 
allowing free movement between P and PS. Basally, where 
the PS is thickest, this fibrous layer conversely becomes so 



A.R. MEAD 



thin on the surface of the P and so intricately associated with 
it, as to be essentially imperceptible. About midway on the P, 
the PS suddenly goes from thick to thin. This creates a 
transverse line of thin folds that incorrectly suggests the PS 
terminates at that level (Mead, 1992, fig. 2). However, when 
there is extreme contraction during preservation, the apical 
edge of the PS actually does pass basally far enough to allow 
the apical structures to elbow out of the PS (Fig. 3). The 
contraction emphasizes the bipartite nature of the P: an 
apical convoluted, transparently ensheathed portion and a 
basal irregularly bulging, opaquely ensheathed portion that 
contains the pilaster (PIL). Internally, the most basal P is 
longitudinally plicate; above that, including the PIL, the 
epithelium is vermiculate-rugate. The PIL is a simple, greatly 
thickened, longitudinal, roundly elevated ridge of the ventral 
penial wall that strongly projects dorsally into the lumen of 
the P. Basally, this ridge terminates into a solid, inverted- 
conical, pendulous verge-like process. Although its margins 
are not well defined, axially the PIL has a more gross 
epithelial texture than the surrounding tissue. The apical vas 
deferens (AVD) is a conspicuously uniformly slender conduit 
(—1.0 mm in width). It lacks the heavy muscular basal 
portion found in C. shuttleworthi, thus the physical support 
for the intromittent organ in C. barriana doubtless is pro- 
vided by the thick, longitudinal P. 

The vagina (V) is a short, nearly uniformly wide conduit, 
about one-third the length of the P. Internally, it is lined with 
vermiculate-rugate epithelium and is without any apparent 
modifications at its junction with the spermathecal duct (SD) 
and free oviduct (FO). The muscular FO is as wide or wider 
than the V, 2-3 times as wide as the SD, and about as long as 
the SD. For their full length, both FO and SD are tightly 
bound to each other by fairly regularly appearing small slips 
of muscle. The junctions of the AVD/FO and spermatheca 
(S)/SD are pulled in close juxtaposition by the tissues of the 
sagittal myoseptum. Just apical to this, the capitate S, about 
the length of the V, is broadly attached to the basal (uterine) 
portion of the spermoviduct. The SD is a thin-walled mostly 
uniformly slender conduit about the caliber of the AVD. Five 
gravid specimens were examined; three with full data had 
been collected near the end of the rainy season in October/ 
November. For such a relatively small species, the eggs are 
quite large (6.8 x 5.4-6.3 x 5.1 mm). Fully gravid specimens 
contained 11-15 eggs, all with heavy, calcareous shells and 
distributed in the full length of the spermoviduct. The ovotes- 
tis acini appear in four or five discrete clusters under the 
columellar surface of the right (apical) lobe of the digestive 
gland. A talon with a round base and an apical, diverticulate 
elongation is present. 

The genital atrium (GA) in this species is unique among 
the achatinids so far dissected. It is comparatively large and 
so shallow that it is essentially a common genital depression, 
immediately within which appear conspicuously the male and 
female orifices. These latter, like twin craters, are individu- 
ally surrounded by low elevated circular walls of smooth 
tissue, which contiguously fuse at their inner margines (Fig. 
3). 

Type material. Sowerby (1890:579) did not designate a 
holotype. The BMNH specimen '89:11.19.2 purchased of 
Sowerby' is here designated the lectotype (Figs. 23, 24; Table 
1). The slightly damaged and trimmed second syntype, 
NMW, 1955:158.832 in the Melvill-Tomlin collection is here 
designated a paralectotype (Oliver, 1983). Remeasurements 



of the lectotype confirm Sowerby's figures except for the shell 
length, which is 41.0 mm rather than '43 mm'. Sowerby's 
illustration is so poorly rendered that it is not precisely 
identifiable with either syntype. 

Type locality. 'Calabar, Africa?' Nigeria, 4° 57' N, 8° 19' 
E. J.C. Reid of the University of Calabar recently confirmed 
this queried locality. Although he has made many excursions 
into the 'relatively undisturbed Oban Hills Forest which 
yields a rich fauna', he found only two (live) specimens along 
a permanent stream at Aking (= Awsawmba) 5° 26' N, 8° 38' 
E, 78 km northeast of Calabar. One of these specimens 
(BMNH) was examined anatomically and conchologically in 
the present study and was found to be typical; the second 
specimen is reportedly in the Tom Pain collection (NMW). 

Distribution. This species has been found essentially along 
the entire expanse of coastal Cameroon from M'Bonge (= 
Bonge) 4° 33' N, 9° 05' E in the north to Itoki 2° 24' N, 9° 50' 
E in the south. Most of the known twenty localities are 
clustered in northwestern Cameroon, spilling over into south- 
eastern Nigeria and extending inland as far as Yaounde 3° 52' 
N, ll°31 o E;Metet3°05'N, 11° 00' E; Ebolowa2° 54' N, 11° 
09' E and Sangmelima 2° 56' N, 11° 59' E. The nine other 
localities are in the environs of Victoria 4° N, 9° E. In all 
localities, seven were shared with C. shuttleworthi and five 
were shared with Leptocala mollicella. Only a single general 
locality record was found for Gabon (Verreaux, 1855 
NHMB) and no record for Equatorial Guinea; but this 
species eventually probably will be found to be limited to the 
northern regions of these two countries. Data labels indicate 
that specimens were collected in plantations in Kumba 4° 38' 
N, 9° 25' E (bananas), Missellele 4° 07' N, 9° 25' E (coca), 
'Buenga' (oil palm), and in primary forests. 



Table 1 


C. barriana - 


Representative shells measurements. 






Greatest Aperture 


Last 


/o 






Whorls 


Length 


Width 


Leng 


h Width whorl LW/L % 


W/L 


6V4 


59.0 


32.4 


36.1 


19.5 


49.0 


82 


55 


Bonge 

(UUZM) 


6V< 


57.5 


30.0 


32.8 


16.9 


46.0 


80 


52 


Victoria 
(ZMB) 


6 


50.7 


27.8 


30.9 


16.8 


41.9 


83 


55 


Idenau 
(SMF) + 


6 


49.0 


29.9 


30.6 


17.5 


40.8 


83 


61 


Bonge 
(SMNH) 


5% 


44.0 


27.4 


29.6 


16.3 


37.8 


86 


62 


Bonge 
(SMNH) 


6 


41.0 


23.3 


24.9 


14.5 


32.3 


79 


57 


Calabar 
(BMNH) 
LectA. 
barriana* 


6 


41.0 


22.8 


26.6 


13.4 


34.0 


83 


56 


Kumba 
(MRAC) 

795.173 


5V2 


36.8 


23.0 


23.6 


13.4 


29.8 


81 


62 


Idenau 
(SMF) 


5 


25.2 


15.3 


16.4 


8.8 


20.6 


82 


61 


Bibundi 
(SMF) 


4% 


20.5 


13.8 


13.2 


7.6 


16.8 


82 


67 


Bibundi 

(SMF) 



Total specimens examined: 125. Sources: BMNH, CMNH, IRSN, MCZ. 
MNHN, MRAC, NHMB, NHMW, NMW, SMF, SMNH, UHZL UUZN, 
ZMB, ZSM. 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



Remarks. This species is commonly encountered in collec- 
tions and often confused with immature Achatina bandeirana 
Morelet, 1866 and A. craveni E.A. Smith, 1881, both of 
which have a proportionately much smaller last whorl. 

Callistoplepa shuttleworthi (Pfeiffer, 1856) 

Figs. 25, 26 

Achatina shuttleworthi 

Pfeiffer, 1856:34, 1859:603, 1868:216, 1877:275. 

Callistoplepa shuttleworthiana 
Ancey, 1888:69. 

Ganomidos shuttleworthi 

d'Ailly, 1896:69, pi. 3, figs. 11-14, text fig. (radula). 

Callistopepla shuttleworthi 

Ancey, 1898:92; Thiele, 1929:560, fig. 644 (radula). 

Callistoplepa shuttleworthi 

Pilsbry, 1904-05:127, pi. 47, figs. 18-20, pg. xv (radula, ex 
d'Ailly); Germain, 1909:90, 1916:248, pi. 10, fig. 4; 
Bequaert & Clench, 1934b:114; Ortiz & Ortiz, 1959:45, pi. 
5, figs. 97, 98, text figs. 28-31 (genit. syst., pallial com- 
plex, jaw, radula); Zilch, 1959:373, fig. 1352. 

Ganomidus shuttleworthi 
Boettger, 1905:170. 

Shell. Shell elongate-ovate, extremely thin, very fragile, 
translucent with a subdued gloss. Whorls 5'/2-5 3 /4, rarely 6, 
noticeably flattened in profile. A somewhat restricting, 
deeply cut second nepionic whorl produces a mammillate 
obtuse apex. The following whorls form a slender conic spire 
as they descend more rapidly than they expand. Sutures 
between nepionic whorls are deep and regular; those between 
postemergent whorls are more shallow and only slightly 
irregular. Last whorl subcarinate, noticeably so in juvenile 
specimens, expanding proportionately, 77% of shell length, 
range for 4'/2— 6 whorls = 73-83% (n = 60). Aperture 
oblique-ovate, external colour pattern sharp and distinct 
from within. Columella usually straight, axial, rarely slightly 
arcuate, inner rim erect with a cord-like thickened crest; 
truncation oblique to very oblique, rarely at right angles. 
Between the third and fourth whorls, the crest of the col- 
umella rolls abaxially on itself to form a hollow tube, there- 
fore an open umbilicus. Between the fourth and fifth whorls, 
this tube narrows and solidifies to form a slender axial cord, 
which is seen in the full grown shell. This series of changes, 
from open to closed, enigmatically has been observed in 
several disparate achatinid species, e.g. A. achatina (Linne, 
1758) and Archachatina spp. Outer lip of shell thin, skewed 
basally, joining the periphery at an acute angle; greatest 
width below midway; this is emphasized by the subcarinate 
nature of the shell. Parietal callus thin, vague. 

Shell ground colour is pale corneous. The first 2'/i whorls 
are unicolorous. Starting near the third whorl, vague round- 
ish, very pale castaneous spots appear both at the suture and 
periphery. At these two levels, the spots quickly assume 
sharply angulate prosocline arrow-shaped patterns, high- 
lighted with a series of parallel transverse elongate white 
flecks. Similar flecks, reminiscent of those in C. barriana, are 
scattered irregularly over the shell above the periphery, 
rarely below. Soon the sutural band fractionates and moves 
increasingly into a subsutural zone. The large arrows at the 
periphery become spirally closely juxtaposed to form an 
essentially continuous dominating colour band. From it, 
slender, nearly parallel light castaneous stripes pass sinuously 



to the subsutural band and transversely to the columella. 

Only rudimentary costae appear in the last part of the 
otherwise smooth first whorl. The entire second whorl is 
conspicuously and uniformly ribbed from suture to suture 
with elevated, deeply cut, nearly orthocline, gross costae, ca 
0.2 mm wide (cf Germain, 1916 pi. 10, fig. 4). In the third 
whorl the now more prosocline costae are soon reduced to 
half their width. At midway in this whorl, an interruption in 
the alignment of the costae marks the end of the nepionic 
whorls. Gradually the costae become wider and finally regain 
their original width in the fifth whorl, only to become 
narrower and somewhat irregular in the last part of the sixth 
whorl. Faint shallow closely spaced spiral lines, starting in the 
second whorl, almost imperceptibly transect the prominent 
costae. There is a delicate, greatly suppressed cancellate- 
granulate sculpture on a vitreous surface below the periphery 
in the upper whorls. This is invaded by the costae in the sixth 
whorl until the entire whorl from suture to columella is nearly 
uniformly costate. No splitting or anastomosis of the costate 
has been observed. 

Soft anatomy. Alcohol preserved specimens available 
12/dissected 5. Cameroon: SMNH 5/2, UUZM 7/3. All 
specimens were collected by Y. Sjostedt. The only two extant 
mature specimens were found by A. Franzen in a medical 
laboratory at UUZM. d'Ailly (1896) reported having access 
to 11 alcohol preserved specimens, which apparently did not 
include Sjostedt's Itoki specimens that were available in the 
present study. 

Body colour as in C. barriana; spade-shaped elevations on 
the posterior foot only slightly less prominent. 

Without having dissected and deciphered first the relatively 
more simple reproductive tract of C. barriana, it would have 
been very difficult to interpret the relationships of the genital 
structure in this species. In essence, the axis of the basal male 
conduit has been greatly foreshortened telescoping the 
homologous structures to such and extent that the pilaster 
(PIL) on the ventral wall of the penis (P) is pivoted 180°, 
forcing the junction of the P and basal vas deferens (BVD) 
deeply into the dorsal aspect of the infolded P, i.e., the upper 
ventral wall of the P and the most basal part of the BVD are 
therefore seen only in the dorsal or lateral views (Figs. 4, 5). 
A dense network of muscle and connective tissue fibrils 
firmly fixes the structures in this permanently partially evagi- 
nated position. This places the aperture of the BVD and the 
contiguous subapical part of the PIL into a basal position 
within the folded penial wall to take the lead in forming the 
intromittent organ at extroversion. The inner smooth shiny 
surface of the extremely thin-walled penis sheath (PS) facili- 
tates seriatim extroversion: PIL-P-PS and finally genital 
atrium, with the BVD and the attenuated penial retractor 
(PR) contained axially within the intromittent organ. Figures 
6, 7, 8 show at progressively deeper levels of dissection these 
relationships from the ventral view. Both PIL and the inner 
penial wall are confluent with a deeply rugate epithelium. It 
should be noted that since the BVD opens directly into the 
lumen of the penial chamber rather than passing through the 
accessory organ to open at its apex, a pilaster rather than a 
verge (penis papilla) is formed. 

The PR is extremely short and, as in the other species of 
Callistoplepinae, it and the BVD are held tightly together by 
the PS apical to the completely enclosed P (Fig. 9). In 
contrast to that in C. barriana, the PR inserts on the right 
columellar retractor (RCR) anterior and strongly ventral to 



10 

the retractor of the right ommatophore (RRO) (Figs. 10, 2). 
Ortiz & Ortiz (1959) missed the diminutive PR in their 
dissections and do not show it in their illustrations. Emerging 
above the PS, the apical vas deferens (AVD) is a large 
muscular thick-walled conduit that in its normal position 
reaches to the peniovaginal angle and doubtless serves both 
as an ejaculatory duct and a physical support for the intromit- 
tent organ. Apical to this, the conduit narrows to half the 
calibre and is thinner walled. 

The vagina (V) is very short, but two to four times wider 
than its length. Near its base, sparse, thin muscle strands 
suggest a primordial vaginal retentor. Internally, the V is 
muscular, thick-walled and longitudinally deeply plicate. 
There is no sharp delineation between it and the broad, 
somewhat thinner walled basal spermathecal duct (SD). This 
latter is so large that it tends to be positioned partly between 
P and V. Both upper SD and free oviduct (FO) are thin- 
walled and of about the same calibre. The clavate spermath- 
eca (S) is broadly attached to the spermoviduct apical to the 
AVD/FO junction. The ovotestis acini are as in C. barriana. 
No specimen was found to be gravid, but a single specimen 
collected in October in Bonge seemed to be near it with a 
very large albumen gland and an inflated spermoviduct. Ortiz 
& Ortiz (1959) examined a single specimen from Fernando 
Poo Island (Macfas Nguema Biyogo) and found the sper- 
moviduct completely crowded with four comparatively large 
white eggs. Seven dried eggs (MCZ no. 219224) measured in 
the present study average 4.7 x 3.7 mm. A diminutive talon 
is present. In contrast to C. barriana, the genital atrium is an 
inconspicuous dimple without superficial embellishments. 

Type material. Pfeiffer (1856) described this species from 
Cuming collection specimens, giving the shell size as 5 1 /: 
whorls and the length-width measurements as 34 x 17 mm; 
later (1859) he gave aperture measurements as 19 x 
11.5 mm. The measurements of the three syntypes in BMNH 
do not match those of Pfeiffer, but are reasonably close. The 
largest syntype has a damaged and repaired last whorl and the 
next largest is atypically slender; therefore the smallest 
specimen (Figs. 25, 26; Table 2) is here selected as the 
lectotype, the other two becoming paralectotypes. No con- 
vincing evidence was found that other syntypes are extant. 
Only four other specimens of the 60 examined in the present 
study exceeded 30 mm in shell length. 

Type locality. The syntypes in the Cuming collection were 
reported to be from ' "Grand Bassam" Africae occidentalis 
(Verreaux)'. All other specimens examined bearing this 
locality were sold by shell dealers, viz. Da Costa, Fulton, 
Geret, Paetel and Preston, who may have taken their cue for 
a locality from the original description. No museum specimen 
has been found with a locality record from the 1400 km 
stretch of continental Africa between Grand Bassam, Ivory 
Coast and the cluster of reliable locality records in northwest 
Cameroon. It is suspected that this is a case of still another 
erroneous Cuming record. Although under the circum- 
stances, we must accept the type locality as 'Grand Bassam', 
it is probable that Cuming's specimens came from Cameroon, 
or perhaps Gabon. The likelihood of an early secondarily 
established population in Grand Bassam prior to 1856 is 
extremely remote, for surely authentic collecting records 
would have appeared in the meantime. Edouard Verreaux (cf 
Crosse & Fischer, 1869) not only collected the syntypes of 
this species, but he also collected the single known specimen 



A.R. MEAD 
Table 2 C. shuttleworthi - Representative shells measurements. 



Greatest Aperture Last % 
Whorls Length Width Length Width whorl LW/L % W/L 



6 


34.0 


18.7 


18.4 


10.3 


26.4 


78 


55 


'G.Bassam' 

(MCZ) 
83441 


6 


32.3 


18.4 


18.4 


10.0 


24.5 


76 


57 


'G.Bassam' 

(BMNH) 

PLec 


6 


31.0 


16.7 


16.4 


9.0 


22.8 


73 


54 


'G.Bassam' 

(BMNH) 

PLec 


5% 


30.9 


18.3 


17.0 


10.7 


24.0 


78 


59 


'G.Bassam' 
(BMNH) 
Lect A . 
shuttle- 
worthi* 


5% 


26.9 


15.8 


14.8 


8.0 


20.7 


77 


59 


Edea 
(MCZ) 


5'/2 


24.1 


14.8 


14.4 


8.0 


19.0 


79 


61 


Gabon 
(MRAC) 
5314 
(Preston) 


5'/: 


22.9 


13.0 


12.1 


6.8 


17.3 


75 


57 


Bibundi 
(SMNH) 


5V< 


20.2 


11.5 


11.8 


6.8 


16.1 


80 


57 


Itoki 
(UUZM) + 


5 


19.4 


11.4 


11.2 


5.9 


15.4 


79 


59 


Bibundi 
(SMF) 


4V2 


16.0 


10.5 


8.9 


5.3 


12.5 


78 


66 


Gabon 
(IRSN) 
(Vignon) 



Total specimens examined: 60. Sources: BMNH. GNM. IRSN. MCZ. MNHN. 
MRAC. NHMW, SMF. SMNH. UMMZ. USNM, UUZM. ZMB. 

of C. barriana from Gabon, now in Bern (NHMB). This 
raises the suggestion that Verreaux, after collecting in Gabon 
and Cameroon, shipped his specimens from Grand Bassam, 
which Cuming assumed was the collecting site. 

Distribution. Leonardo Fea was the first to discover this 
species on Fernando Poo Island (= Macias Nguema Biyogo) 
of Equatorial Guinea 3° 30' N, 8° 40' E (Germain, 1916:249. 
Subsequently, Ortiz & Ortiz (1959:45) reported it from Basile 
and Mongola on that island. Nine reliable localities on the 
mainland in Cameroon define a limited coastal belt, ca 280 x 
120 km with N'dian 4° 55' N, 8° 53' E in the north; Metet 3° 
05' N, 11° 00' E in the east; and Itoki 2° 24' N, 9° 50' E in the 
south. Other Cameroon localities: Albrechts Hohe 4° 38' N, 
9° 25' E; Mukonje (= Mukonye) 4° 37' N 9° 30' E; Bibundi 4° 
13' N, 8° 59' E; Edea 3° 48' N, 10° 08' E; Lokoundje 3° 13' N, 
9° 55' E. A specific locality record for Gabon was not found, 
but Vignon, through Ancey (1888:69), reports them as rare in 
Gabon at the edge of forest streams. They probably do not 
extend south of the Ogooue River. 

Callistoplepa tiara Preston, 1909 - A Misidentification 

Preston (1909:183, pi. vii, fig. 9) described Callistoplepa tiara 
from 'Bitze [= Bitye], near the River Ja [Dja], Cameroons' 
(3° 01' S, 12° 22' E). He indicated neither the collector nor 
the number of specimens he had; however, some specimen 
labels (BMNH, MRAC) specify that G.L. Bates was the 
collector. Between 1908 and 1912, Preston distributed ten 
known syntypes, each bearing the full locality information 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



11 



RCR 






RCR 






Fig. 9 C. shuttleworthi , basal genital structures (UUZM). 

Fig. 10 C. shuttleworthi, right branch of columellar muscle showing anterioventral attachment of the penial retractor. 

Fig. 11 Leptocala mollicella, basal genital structures (MRAC no. 796.850). 

Fig. 12 L. mollicella, penis sheath, penial retractor and penial wall cut longitudinally and spread to reveal the pendulous pilaster within 

(MRAC no. 795.638). 
Fig. 13 Same, cutaway of pilaster to show basal vas deferens joining the penial sacculus, which leads to the aperture of the pilaster. 
Fig. 14 L. petitia, basal genital structures (MRAC no. 214.044). 



12 



A.R. MEAD 



and Preston as the source. These syntypes are currently to be 
found in the following museums: BMNH (3: 
no. 1908. 7. 1.13-14, and MacAndrew Coll.), NMW (Melvill- 
Tomlin Coll. no.1955. 158.826; Oliver, 1983:1), IRSN 
(Dautzenberg Coll. no. 169), ZMB (no.62345), MNHN, 
MRAC (no.5760), RMNH, UMMZ (Bryant Walker Coll. 
no. 142031). These vary in size from 6V2, 63.0 x 31.3 to 5 l h, 
44.2 x 25.7. Preston probably placed his presumed new 
species in Callistoplepa because of the very thin shell, the 
Cameroon type locality, and the fact that the size, general 
shape and peripheral colour pattern of his specimens were 
reminiscent of C. barriana. However, upon examination of 
the shell sculpture in the present study, all syntypes were 
found to be juvenile Achatina bandeirana Morelet, 1860. 

In Cameroon, A. bandeirana and the closely related A. 
iostoma Pfeiffer, 1854 and A. balteata Reeve, 1849 are 
sympatric and it is not uncommon to find mixed lots of these 
three species in museum collections. Preston, himself, appar- 
ently had a mixed lot from which his syntypes were selected. 
He sent a 'cotype' of Callistoplepa tiara to Dupuis (IRSN, 
General Coll.); however, its locality record was simply 'Cam- 
eroon'. After Dupuis (1923) examined this specimen, he 
concluded that it probably was a juvenile A iostoma. In 1934, 
Bequaert also saw this specimen and confirmed Dupuis' 
conclusion. Their identifications were corroborated in the 
present study because this 'cotype' specimen revealed the 
following characters in contrast to those of the syntypes 
identified as A. bandeirana: 1) upper whorls not convex, but 
form a nearly straight-sided pyramid; 2) apex more acute 
rather than blunt; 3) a slight but apparent peripheral carina is 
present in the early whorls; and 4) sculpture is formed by 
finer, more uniform, elevated beads that do not evolve into 
minute prosocline arcuate welts in the sixth to seventh whorls 
(cf Bequaert & Clench 1934a fig. 3). This last character is 
diagnostic for A. bandeirana; but it is inadequately developed 
in the very immature specimen of five to six whorls, thus such 
individuals of the three species may appear to be alike. 

Dupuis' unique 'cotype' persuaded Bequaert to assume 
that all syntypes of C. tiara were juvenile A. iostoma and he 
so identified them in collections (BMNH, IRSN, ZMB, 
RMNH) and in his publications (Bequaert, 1950:39; B. & 
Clench 1934a:13; 1934c:114). Dautzenberg was similarly 
impressed and was moved to place with his 'cotype no. 169' an 
added notation, 'Erreur de Preston, C'est un jeune Achatina 
iostoma Pfeiffer'. This was unfortunate because Dautzen- 
berg's specimen, with full C. tiara field data, is shown now to 
be an immature A. bandeirana. IRSN thus has a true syntype 
in the Dautzenberg Collection and questionable 'cotype' in 
the General Collection, which latter is here confirmed to be 
A. iostoma and not a bona fide syntype. A somewhat similar 
situation exists at BMNH, which has three valid syntypes. A 
fourth specimen in the Connolly Collection (BMNH 
no. 1937: 12.30.3684) was sent by Preston and labelled 'Callis- 
toplepa tiara Pr.' (apparently in his writing) but without any 
locality data, except 'Bitz' in the accession book. Connolly 
had his doubts about the identification and relabelled it 
'Achatina ? balteata Rve juv.' Bequaert also saw it in 1933 
and referred to it as A. iostoma. This now proves to be still 
another juvenile A. bandeirana and is here considered a 
doubtful eleventh syntype of C. tiara. 

It should be noted that A. bandeirana is a wide spread, 
highly variable Lower Guinea species complex involving A. 
b. arenaria Crowley & Pain, 1961; A. b. mayumbensis C. & 
P., 1961; A. paivaeana Morelet, 1866, (1868); and A. dohrni- 



ana Pfeiffer, 1870. It is found from Cameroon to northern 
Angola (7° N-10° S) and fans north and east into Gabon, 
Central African Republic, Congo Republic and Zaire. A 
study of this complex is in progress. 

Preston did not designate a type, but he retained in his own 
collection the specimen that was illustrated in his description 
of this species. This syntype is here selected as the lectotype 
(measurements: 6; 49.4 x 26.7; aperture 30.9 x 13.8; last 
whorl 40.0 mm). It is now in Tervuren (MRAC no.5760) and 
can be precisely identified by the unique configuration of the 
map-like pattern on the last whorl. This pattern is caused by 
the irregular lifting up of the thin outer periostracal layer 
from the durable inner periostracal layer, allowing an air 
space between. This produces blotchy grey-white patches, 
which probably provide cryptic coloration. The juvenile and 
mature specimens of both A. bandeirana and A. iostoma 
commonly have these patches, which have been referred to as 
'hydrophanous streaks' (Bequaert & Clench, 1934a: 15). They 
apparently are homologous to the conspicuous white flecks 
on the shells of Callistoplepa barriana and C. shuttleworthi 
and may have contributed to Preston's decision to put his 
species in this genus. 

Leptocala 

Petitia 

Jousseaume, 1884:171 (non Chitty, 1857); d'Ailly, 

1896:71; Bequaert, 1950:138 (type species: Petitia petitia 

Jousseaune, 1884). 
Leptocala 

Ancey, 1888:70, 1898:92 (type species: Achatina mollicella 

Morelet, 1860); Thiele, 1929:560; Bequaert & Clench, 

1934c: 116; Ortiz & Ortiz, 1959:24. 
Achatina (Leptocala) 

Pilsbry, 1904:72; Spence, 1928:213; Bequaert, 1950:138; 

Zilch, 1959:366; Vaught, 1988:89. 
Achatina (Leptocola) 

Kobelt, 1910:66 (non Gerstaecker, 1883). 
Leptocala (Leptocala) 

Bequaert & Clench 1934b:272. 

Pilsbry (1904:73, 75) reduced genus Leptocala to subgeneric 
rank in Achatina and placed within it his new Section 
Leptocallista. Thiele (1929:560) returned Ancey's Leptocala 
to generic rank and retained within it Sections Leptocala and 
Leptocallista. Bequaert & Clench (1934b:274) elevated these 
sections to genus and subgenus, respectively. In 1950, 
Bequaert placed both names as subgenera of Achatina. Zilch 
(1959:366) followed suite. The present studies of the soft 
anatomies demonstrate that these two genus-group taxa are 
in separate subfamilies because the East African Leptocallista 
is anatomically allied to Lissachatina and therefore is an 
achatinine. 

Bequaert & Clench (1934b, c) announced that the Cam- 
eroonian Pseudoglessula efulensis Preston, 1908 might belong 
to Leptocala and stated that the type could not be located in 
the British Museum. The holotype (no. 5309) and the 
paratype (no. 97435) of this species were found during the 
present study in Tervuren (MRAC) and clearly proved to 
belong to the Subulinidae. Ancey (1888:71) incorrectly 
placed Achatina polychroa Morelet, 1866 in Leptocala; 
Bequaert (1950:48) believed it belongs in subgenus Pintoa of 
Achatina. A final decision depends upon a study of its soft 
anatomy. 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



13 



The ancestral stock of the two closely related, remaining 
species in this genus, L. mollicella and L. petitia, probably 
became separated in fairly recent times by a vicariance event 
- possibly the development of the Ogooue River. 

Because of the unique microsculpture and the somewhat 
smaller shell aperture, Leptocala up until now has escaped 
suspicion of being closely related to Callistoplepa. The genus 
is limited to the southwestern portion of Lower Guinea from 
northwestern Cameroon to far western Zaire. 

Key to Species 

Shell 6-6V2 whorls; spire conic; exceedingly fine distinct 
vertical and spiral lines form shallow minute engraved rhom- 
boids. Pilaster verge-like, cylindrical, vertically suspended 
from the apex of a dome-shaped penis; basal vas deferens 
obscured by penial retractor. North of Ogooue River in 
Gabon, Equatorial Guinea, Cameroon and probably south- 
eastern Nigeria mollicella 

Shell 6V2-7 whorls; spire slender conic; exceedingly fine 
closely appressed vertical vermiculate-granulate sculpture 
obliterates the spiral lines, especially on the upper whorls. 
Pilaster potato shaped, somewhat compressed, attached for 
nearly its full length along a diagonal right ventrolateral axis 
of a hull shaped penis; basal vas deferens conspicuous in 
ventral view. South of Ogooue River in Gabon, Congo 
Republic, western Zaire and probably Cabinda, Angola- 
petitia 

Leptocala mollicella (Morelet, 1860) 

Figs. 27, 28 

Achatina mollicella 

Morelet, 1860:189; Pfeiffer, 1868:216; 1877:275; Vignon 

(in Ancey, 1888:70); Pilsbry, 1904:29. 
Achatina pulchella 

von Martens, 1876:258, pi. 3, figs. 1, 2 (syntype) {non Spix 

& Wegner, 1827; non Pfeiffer, 1857); Ancey, 1888:70; 

Pilsbry, 1904:73, pi. 34, fig. 14 (ex von Martens). 
Leptocala mollicella 

Ancey, 1888:70, 1898:92; Thiele, 1929:560; Bequaert & 

Clench, 1934b:273. 
Achatina smithi 

Sowerby, 1890:579, pi. 56, fig. 3 (holotype, monotypy; 

non Craven, 1880). 
Achatina sowerbyi 

E.A. Smith, 1890:392 (new name for A. smithi). 
Petitia pulchella 

d'Ailly, 1896:71; Boettger, 1905:170. 
Achatina (Leptocala) mollicella 

Pilsbry, 1904:73; Spence, 1928:213, pi. 2, fig. 5; Bequaert, 

1950:138; Zilch, 1959:366, fig. 1342. 
Achatina (Leptocala) pulchella 

Germain, 1916:154, 241, pi. 6. figs. 11, 12. 
Leptocala (Leptocala) mollicella 

Bequaert & Clench, 1934b:273. 
Leptocala mollicella zenkeri 

Bequaert & Clench, 1934c: 118, pi. 1. figs. 5-7, pi. 2, fig. 

13 (holotype, 3 paratypes). 
Leptocala mollicello zenkeri 

Bequaert & Clench, 1934c:119 (lapsus calami). 
Achatina (Leptocala) mollicella petitia 

Bequaert, 1950, pi. 58, fig. 4. 
Achatina (Leptocala) mollicella zenkeri 



Bequaert, 1950:138. 
Leptocala pulchella 

Ortiz & Ortiz, 1959:25, pi. 5, fig. 99. 

Shell. Shell obovate, glossy, translucent, thin but sturdy; 
periostracum tenaceous. Whorls 6-6V2, moderately convex. 
Spire conic; apex broadly obtuse; sutures moderately deep, 
fine, straight or slightly irregular. Last whorl expanding at a 
somewhat greater rate than the upper whorls, 72% of shell 
length, range for 4 3 /4-6>/2 whorls = 69-75% (n=34). Aperture 
elongate inverted ear-shape, pale milky within. Columella 
short, variably straight to slightly arcuate and twisted, trans- 
versely to obliquely truncate, basal crest slightly elevated in 
juvenile specimens. Outer lip thin, joining the periphery at a 
broadly acute angle. Parietal callus minutely granular, shiny, 
concolorous. 

Shell ground colour is pale fulvous, rarely somewhat 
darker. Most specimens have a distinct but subdued pattern 
of pale yellow-brown, slender, strongly parallel, nearly 
straight or somewhat sinuous stripes, usually 0.2-0.3 mm 
wide, alternating with ground colour bands of about the same 
width. The banding may be slightly coarser and more con- 
spicuous in the fifth whorl and above. Often apparently 
unicolorous or weathered specimens under proper lighting 
and magnification will be seen to have this characteristic 
pattern at least in limited areas. This is witnessed in Bequaert 
& Clench's (1934c) figure 7 of their Leptocala mollicella 
zenkeri, which they report is 'without any darker markings'. 

The first I-IV2 whorls are smooth and very shiny. Short 
vertical or arcuate lines begin to appear in the second whorl, 
often concentrated at the suture below. These soon elongate 
into delicate, narrow, closely packed vertical lines that span 
the full width of the whorl. At 2 ] k whorls, there is a 
conspicuous diagonal demarkation between the nepionic and 
the postemergent whorls. At this demarkation, spiral lines 
that immediately previous to this were sparce, short and 
ghost-like, quickly form 35^40 nearly evenly spaced exceed- 
ingly shallow, but sharply engraved lines. These lines cross 
the vertical lines and cut the surface into minute engraved 
rhomboids (see Bequaert & Clench, 1934c fig. 13). In the 
following whorls, these spiral lines become more numerous 
and somewhat wavy, suggesting the surface had been evenly 
and shallowly combed. Beginning at 2'/2 whorls, subtile 
prosocline, more sparce growth wrinkles compete with the 
vertical lines; these may impact the suture directly or arcu- 
ately. There is little reduction in the intensity of this engraved 
pattern below the periphery, although there is a slight reduc- 
tion in caliber. In the last half of the last whorl, the rhomboid 
pattern may essentially disappear, leaving the growth 
wrinkles to dominate. Throughout, the shell characteristically 
remains remarkably smooth and shiny. A subcarina is present 
in the early whorls, but this disappears in the fifth whorl. 

Soft anatomy. Alcohol-preserved specimens available 
12/dissected 4. Cameroon: MRAC 3/3, UHZI 1/0; ZMB 7/0; 
ZMUC 1/1. 

Body colour of preserved specimens, including the head 
and anterior edge of the mantle, is pale-cream fuscous. 
Dorsally, there is a diffuse grey wash that shades darker 
anteriorly. A 1-2 mm wide dark gray band with diffuse 
borders, appears immediately behind the ommatophores and 
extends posteriorly along the middorsal neck to the edge of 
the mantle. Surface of body reticulate-microtuberculate. Foot 
without structural elaborations. 



14 



A.R. MEAD 



The most prominent feature of the genital system (Fig. 11) 
is the massive, muscular basal female conduit that seems to 
be an ill defined fusion of vagina (V), spermathecal duct (SD) 
and free oviduct (FO). Sessile to this, obliterating the penio- 
vaginal angle, is the contrastingly short, broad penis sheath 
(PS). Crowded together and projecting from the apical collar 
of the PS are the long apical vas deferens (AVD) and the very 
short penial retractor (PR). In fact, this latter is so short that 
only its insertion on the broad right columellar retractor 
(RCR) may be seen. Like that of L. petitia and Callistoplepa 
barriana, it inserts far posterior on the RCR (Fig. 2). The 
tripartite AVD starts as a slender tube, but soon enlarges to 
form the thick- walled ejaculatory duct. This in turn narrows 
and then once again enlarges into a thin-walled conduit 
(possibly a secondary seminal vesicle) before fusing with the 
FO to form the spermoviduct. When the PS is cut longitudi- 
nally, it is found to be extremely thin and attached basally 
only about half way down on the stubby, dome-shaped penis 
(P). Both the inner surface of the PS and the outer surface of 
the P are smooth and shiny, facilitating extroversion. In this 
aspect, the PR is seen to hood over the apical P and, about 
one-quarter the way down, blend with the substance of the 
thick-walled P. When the PR is split longitudinally and its 
muscle bands spread apart (Fig. 12), the basal vas deferens 
(BVD) is seen within, discreet and without muscle or connec- 
tive tissue attachment. Vertically cutting the penial wall 
reveals a conspicuous, pendulous, vertically oriented pilaster 
(PIL) whose thick transverse-diagonally textured brownish, 
glandular epithelium is continuous with the inner wall of the 
penis. Ventrolaterally on the PIL is a 1 mm vertical eccentric 
apertural slit. This leads internally to a funnel shaped penial 
sacculus, which joins the extension of the BVD in a dense 
mass of connective tissue. It is clear at this point that the PIL 
has been formed by a permanent partial eversion of the P. In 
the process, the basal-most BVD, with its diagonal anchoring 
muscle strands, forms the axial conduit of the PIL (Fig. 13). 
During extroversion, to form the intromittent organ, the PIL 
would take the lead, followed by the basal P, and finally the 
PS, which would contain the BVD, PR, and a portion of the 
RCR. Both PR and RCR would be involved in the introver- 
sion process. 

The alignment of the V, SD and FO insures that the 
intromittent organ will be channeled directly to the SD. This 
has been accomplished not only by a large knob of tissue 
eccentrically blocking the narrow lumen of the FO, but also 
by a massive buildup of muscular tissue surrounding the large 
lumen of the basal SD. These modifications, in turn, tend to 
wedge the SD between P and FO (Fig. 11). Internally, there 
is a sharp division between the thick-walled, heavily muscular 
V, with its many narrow, tightly compressed, vertical plicae, 
and the thick-walled basal half of the SD (functionally an 
extension of the V), with its several bold, deep, coarse 
vertical plicae. Cutting across these latter plicae are trans- 
verse vermiculate rugae that produce a grossly serrate texture 
on the crests of the plicae. This rough texture seems to 
complement that of the P and PIL. All these structures 
obviate the necessity of a distinct muscular vaginal retentor 
found in many species of Achatininae. The apical SD is 
thin-walled, as is the clavate spermatheca (S). The apical 
saccular FO probably serves to hold the large egg immedi- 
ately before expulsion; basally, however, it is thick-walled, 
with a narrow lumen. An 'elbow' tends to form at the 
junction of the two parts. The lower portion and the collar- 
like thick-walled V doubtless serve as an ovijector. 



Three of the four dissected specimens had the uterus and 
oviduct crowded with 4, 6 or 8 relatively large, off-white 
hard-shelled eggs, measuring 4.3 x 3.7-5.1 x 4.4 mm. Their 
long dimension is ca. 15% of the adult shell length, which is in 
strong contrast, for example, to that relationship in Achatina 
achatina at ca. 5%. All gravid specimens were collected 
September-November, just before the long dry season, and 
because of the demands of producing eggs, they manifested 
considerable emaciation, especially in the digestive system. 
Such observations raise the unanswered question of longevity 
in this small species. In a single specimen, six ovotestis acini 
were found under the columellar surface of the apical lobe of 
the digestive gland. The talon is extraordinarily long and 
slender and without a basal enlargement. 

Type material. The type of Morlet's Achatina mollicella, 
collected by Vignon, has never been illustrated and its very 
existence has been uncertain. It is not in the Morelet material 
in the Paris, Geneva or Tervuren collections. Fulton (1920) 
reported that he had purchased the Morelet land and fresh- 
water shells in 1892, including 'all the types', but that in 
transit between Dijon and London many of the fragile 
specimens, including some types, were broken. The BMNH 
accession book under date 2 April 1893 confirms this infor- 
mation. It lists the accessioned types (pp. 230-254, 2049 
entries) including a single entry indicating that there were 
only two specimens: '93.2.4.119-120 Achatina mollecella, 
Gabon'. However, neither of these two syntypes have the 
length-width measurements of '18 x 12 mm' given by More- 
let (1860) for an individual 6 whorl specimen. Under the 
circumstances, it is most likely that the 6 whorl syntype is the 
one Morelet had in hand. Further, there clearly is an error in 
his reported measurements because in the 35 specimens 
examined in the present study, the shell width averages 52% 
of shell length, not 67% as would be the case if Morelet's 
measurements were correct. For these reasons, the larger 
syntype BMNH no.93.2.4.119 is here selected as the lecto- 
type (Figs. 27, 28; Table 3) and BMNH no.93.2.4.120 
selected as paralectotype. Morelet's incorrect measurements 
have contributed greatly to the confused synonymies of his 
valid species and Jousseaume's valid species Petitia petitia. 

The lectotype and 10 paralectotypes of von Martens' junior 
subjective synonym Achatina pulchella are in Berlin (ZMB; 
Kilias, 1992), type locality Bonjongo, Cameroon. Additional 
single specimens here designated as paralectotypes have been 
found and labeled in Stockholm (SMNH no. 4282) and Ter- 
vuren (MRAC no. 5315). The holotype (monotypy) of Sower- 
by's junior subjective synonym and junior primary homonym 
Achatina smithi no. 89. 11. 18.1 is in London (BMNH), type 
locality 'Calabar, Africa?' The holotype of Bequaert & 
Clench's Leptocala mollicella zenkeri (1934c fig. 6) plus two 
paratypes (their fig. 7 and one unfigured) are in Berlin 
(ZMB; Kilias, 1992). A third paratype is at Harvard (MCZ) 
under No. 98687, which was identified by the distinctive mark 
at the junction of the ultimate and penultimate whorls (cf. 
their fig. 5). All these were from Yaounde. A fourth 
paratype, unfigured but listed on their page 119 is from Bitye 
(BMNH, no.1908.6.3.2; Table 3). 

Type locality. Morelet (1860) lists it as, 'Habitat, rara, in 
sylvis Guinea.' His two syntypes in BMNH were more 
specifically labeled 'Gabon,' which is included in the early 
broad generic geographic term 'Guinea.' 

Distribution. Sowerby (1890) described Achatina barriana 



NEW SUBFAMILY AND GENUS ACHATINIDAE 
Table 3 L. mollicella - Representative shells measurements. 







Greates 


Aperture 


Last % 






Whorls 


Length 


Width 


Length 


Width whorl LW/L % 


W/L 


6'/4 


39.8 


18.7 


17.5 


9.3 


27.7 70 


47 


Bitya 
(BMNH) 
Para L. m. 
zenkeri 
1908.6.3.2 


6'/2 


37.5 


17.7 


17.3 


8.9 


26.5 71 


47 


Ebalowa 
(UMMZ) 


6»A 


35.8 


17.5 


18.4 


8.7 


26.5 74 


49 


Yaounde 
(UMMZ) 


6V2 


33.3 


17.3 


15.9 


10.3 


23.4 70 


52 


Olounou 
(MRAC) 
796.850 


6 


30.5 


16.1 


15.6 


8.2 


22.3 73 


53 


Kribi 

(MRAC) 

795.638 + 


6 


30.0 


14.8 


15.0 


7.5 


21.4 71 


50 


Calabar 
(BMNH) 
Holo/L 
s mil hi 


6 


26.7 


13.4 


12.8 


6.2 


18.9 71 


50 


Gabon 
(BMNH) 
Lect. A. 
mollicella 

93.2.4.119 


6 


26.6 


14.0 


13.6 


6.9 


19.1 72 


53 


Bonjongo 
(MRAC) 
5315 PLec 
A. 
pulchella 


5'/2 


23.6 


13.1 


12.4 


6.6 


17.3 73 


55 


Gabon 
(BMNH) 
PLec A. 
mollicella 
93.2.4.120 


4% 


16.6 


10.4 


9.1 


5.0 


12.2 73 


63 


Nyong 
(ZMUC) + 



Total specimens examined: 35. Sources: BMNH. IRSN. MNHN. MRAC. 
NHMW. NMW, SMNH. UMMZ. ZMUC. 



and A. smithi (= L. mollicella) at the same time and indicated 
for both that the locality was 'Calabar, Africa?' (4° 57' N, 8° 
19' E). Both specimens were from the Cuming collection, in 
which a number of other locality records from time to time 
have been questioned or proven incorrect. J.C. Reid of the 
University of Calabar has collected near Calabar what is now 
confirmed as Callistoplepa barriana, but so far no L. molli- 
cella. It is altogether possible that this latter species eventu- 
ally will be found in Nigeria because Mbonge, Cameroon, a 
known endemic locality for this species, is only ca. 70 km to 
the southeast in a similar environment. Ortiz & Ortiz 
(1959:26) have reported the western-most records for this 
species from four localities on Fernando Poo Island ( = 
Macias Nguema Biyogo) of Equatorial Guinea 3° 30' N, 8° 
40' E. Seventeen locality records on the continent cluster in 
the northwestern corner of Cameroon, with the extremes 
being Mbonge 4° 33' N, 9° 05' E in the North, Molobo 4° 01' 
N, 14° 19' E in the East, and Efulen 2° 42' N, 10° 30' E in the 
South. Vignon through Ancey (1888:70) records this species 
as being very rare in the forests of Gabon. It probably is not 
found south of the Ogooue River. 
In the specimens examined, there was a high direct correla- 



15 

tion between greater shell size and distance from the sea- 
coast, e.g. the largest specimen seen in this study is from 
Bitya on the river Dja, ca. 260 km from the coast 3° 01' N, 12° 
22' E (BMNH no. 1908.6.3.2; Table 3). 

Leptocala petitia (Jousseaume, 1884) 

Figs. 29, 30 

Petitia petitia 

Jousseaume, 1884:172, pi. 4, fig. 4a, holotype, monotypy 

(non Chitty, 1857); Bequaert, 1950:138. 
Achatina (Leptocala) mollicella petitia 

Pilsbry, 1904:73, pi. 34, fig. 15 (ex Jousseaume, 1884); 

Bequaert, 1950:138, pi. 58, fig. 4. 
Leptocala mollicella petitia 

Bequaert & Clench, 1934b:273. 

Shell. Shell ovate-elongate, thin but not fragile; last whorl 
shiny, upper whorls less so. Whorls 6V2— 7, moderately con- 
vex. Spire slender conic; apex narrowly obtuse; sutures fine, 
almost without irregularities. Last whorl expanding propor- 
tionately to upper whorls, 70% of shell length (n=7); fourth 
and fifth whorls subcarinate. Aperture elongate-oval, milky 
within. Columella short, straight, transversely to obliquely 
truncate. Outer lip thin, arcuate, joining the periphery at an 
acute angle. Parietal callus minutely granular, shiny, concol- 
orous. 

Shell ground colour is pale corneous. At the junction of the 
fifth and sixth whorls, diffuse yellow-brown stripes 
(0.3-0.5 mm wide) alternate with wide ground colour bands 
(0.5-0.7 mm); these are approximately the same width on the 
early whorls, but become slightly or much narrower, more 
distinct and closer together on the last whorl, or nearly 
disappear; they may be variously straight, diagonal or 
rippled. 

The last quarter of the first whorl has nearly imperceptible 
surface irregularities that originate close to the suture, where 
they evolve into a series of closely packed crescentic lines. 
They quickly multiply axially into five or six horizontal series 
of short crescentic lines. These gradually fuse vertically to 
form very narrow, crowded, thread-like, prosocline welts. A 
fairly conspicuous diagonal line, near mid third whorl, marks 
the end of the nepionic whorls. Near there, the welts become 
superficially engraved with a vertically oriented, exceedingly 
fine vermiculate-granulate sculpture, which is reminiscent of 
the much coarser sculpture of Achatina (Tripachatina) 
vignoniana Morelet, 1874. Gradually, the welts diminish and 
the more sparce growth lines emerge, leaving the rash-like 
microscopic sculpture to dominate. This is best seen in 
subdued light at a low angle. The sculpture may diminish and 
become more sparce between the third and fourth whorls, as 
it does in the holotype, or it may continue at essentially the 
same caliber until the fourth or fifth whorl. At a certain point 
in the diminution, and if the light intensity is properly 
adjusted, ghost-like, engraved spiral lines, here and there, 
spaced as in L. mollicella, can be distinguished, especially 
below the periphery, where the sculpture is somewhat 
reduced in calibre. The sculpture may extensively obscure 
these spiral lines and all but traces of a rhomboid pattern, or 
it may become so sparce on the lower whorls as to allow the 
sharp spiral lines to dominate. It is almost as if the 
vermiculate-granulate sculpture were superimposed upon the 
typical sculpture of L. mollicella in a variably decreasing 
intensity from apex to base. As a result, the surface of the 



16 



A.R. MEAD 




Fig. 15 L. petitia, penis sheath and penial wall cut diagonally in right ventrolateral view and spread to expose the pilaster. The cutaway 

shows the basal vas deferens joining the aperture of the pilaster (MRAC no. 214.044 & 212.583). 
Fig. 17 Bequaertina pintoi, basal genital structures (NM). 
Fig. 18 B. pintoi, penis sheath cut and spread laterally. Conduits transected. Most of the obscuring basal eversion muscle bands have been 

removed. 
Fig. 19 B. pintoi, hermaphroditic duct system. 

Fig. 20 Bequaertina graueri, basal genital structures (MRAC no. 610.342 & 610.303). 
Fig. 21 B. graueri, penis sheath cut and spread. 
Fig. 22 B. graueri, frontal plane through basal male and female conduits. Bar scale = ~ 1 mm. A.R.M. del. 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



17 



shell has less luster than in this latter species. This peculiar 
microscopic sculpture on the upper whorls is determinative. 

Soft anatomy. Alcohol preserved specimens available 
2/dissected 2. Congo Republic: MRAC 1/1; Zaire: MRAC 
1/1. These apparently are the only such specimens extant. 
Both had small body masses and were withdrawn far into 
their thin shells because they were collected during the dry 
season and were inadequately drowned before preservation. 
However, most of the soft parts were successfully extracted 
with only minimal damage to one shell. 

Body colour and texture as in L. mollicella. 

Upon exposing the reproductive tract (Fig. 14), the most 
noticeable anatomical feature is that both the unusual hull- 
shaped penis (P) and the large basal vas deferens (BVD) 
show through the thin, nearly transparent, but substantial, 
penis sheath (PS). Typical of the Callistoplepinae, the PS also 
enshrouds the very short penial retractor (PR). As in L. 
mollicella and Callistoplepa barriana, this latter inserts far 
posterior on the large right columellar retractor (RCR). Also 
conspicuous is the apparently inordinately long bipartite 
apical vas deferens (AVD), with a nearly evenly broad 
muscular basal portion and a thin-walled, somewhat undulant 
apical portion. When the PS was cut vertically and the edges 
pulled laterally, it was found to extend essentially to the base 
of the P. In the first dissected specimen, from Lukula, Zaire, 
the exposed, large BVD appeared out of proportion and 
excessively deeply wedged into the ventral surface of the P. 
Similarly, the navicular P, with its diagonal left ventrolateral 
orientation, seemed enigmatically distorted. But the second 
specimen, from Kayes, Congo Republic, ca. 180 km to the 
north, had almost identical proportions and alignment, thus 
essentially removing the suspicion that there had been exces- 
sive distortion. The relatively thin penial wall of the first 
specimen was cut along a midventral, vertical line. Immedi- 
ately below the surface was a large, obstructing mass of penial 
wall tissue whose angulate orientation could not safely be 
explored. Consequently, in the second specimen, a diagonal 
cut was made along the long axis of the oblong P. This 
revealed in right ventrolateral aspect a comparatively huge, 
somewhat compressed potato-shaped pilaster (PIL) attached 
diagonally along nearly its full length ventrolaterally on the 
inner basal penial wall, parallel to the adjacent crowded BVD 
(Fig. 15). In essence, the wall of the basal half of the P was 
hardly more than a thin, tight-fitting cover for the PIL. The 
surface of the PIL and the inner wall of the P, similar to that 
of L. mollicella, was covered with transverse, diagonal, 
anastomosing rugae. Irregularities in the rugae revealed a 
small basal aperture. Cutting basally into the 2.6 mm PIL 
exposed the short basal-most BVD narrowing rapidly 
through dense connective tissue to connect with this aperture. 
Apically, the PIL is a solid mass of penial wall tissue. 
Collectively, the relationships in these structures are reminis- 
cent of those in C. shuttleworthi , particularly with respect to 
the exposed BVD pushing ventrally far down into the par- 
tially evaginated P (Figs. 4, 5). 

In this species the basal female conduit, externally and 
internally, is much less gross than in L. mollicella. The vagina 
(V) is a distinguishable, more slender porion of the conduit. 
Similarly, the basal spermathecal duct (SD) is less muscular 
and tends less to interject itself between PS and the free 
oviduct (FO). However, the FO, muscular at the base and 
thin-walled apically, is less robust yet comparatively more 
prominent in this species. Although shown spread apart in 



Fig. 14 for clarity, the FO and basal SD are actually held 
tightly together by many small, short muscle bands, probably 
providing support for the SD at termination of copulation. In 
that natural position, the clavate spermatheca (S) is attached 
by thin muscle bands and connective tissue to the uterine 
portion of the spermoviduct, well above the junction of AVD 
and FO. The eggs are not known but are probably on a par 
with those of L. mollicella. 

Type material. The holotype (monotypy) (Figs. 29, 30; 
Table 4) in the Jousseaume collection in Paris (MNHN) was 
collected by L. Petit. 

Type locality. At the mouth of the River N'toc, which 
disappears in the Mayumba Lagoon, Gabon 3° 25' S, 10° 39' 
E. 

Distribution. Gabon: type locality. Congo Republic: Sibiti 
3° 41' S, 13° 21' E (SMNH), Kola 4° 03' S 1 1° 44' E (MRAC), 
Kayes 4° 26' S, 11° 23' E (MRAC). Zaire: Lukula 5° 21' S, 13° 
02' E (MRAC). All known localities are south of the Ogooue 
River of Gabon. This species will probably be found in 
Cabinda, Angola. 

Remarks. In addition to the holotype in Paris (MNHN), 
there are only six known specimens of this species, five in 
Tervuren (MRAC), collected by Dartevelle, and a single 
specimen in Stockholm (SMNH). The explanation for its 
apparent rarity probably rests in the fact that there has been 
much less professional collecting in south coastal Gabon and 
the Congo Republic than in Cameroon, where L. mollicella is 
not a rarity. In this limited number of specimens extant, there 
is a north to south gradient of more intense vermiculate- 
granulate sculpture and reduced rhomboid pattern. If the 
substantive differences in the soft anatomies had not been 
known, this taxon might well have been assumed to be no 



^/UGANDA ,, 










ZIMBABWE / 

■ 

29.) 




Fig. 16 Distribution of Bequaertina. O = Bequaertina fraterculus , 
% = B. graueri, □ = B. pellucida, ■ = B. pintoi, A = B. marteli. 
Where possible, all localities were checked with the volumes of 
the U.S. Board on Geographic Names. 



18 



A.R. MEAD 



Table 4 L. petitia - Representative shells measurements. 

Greatest Aperture Last % 

Whorls Length Width Length Width whorl LW/L % W/L 



7 32.5 14.8 

6V2 30.9 14.2 

6'A 30.8 14.8 

6V2 29.0 13.7 

6 28.6 14.6 

6 23.9 12.3 

5 20.8 13.3 



14.3 7.5 21.5 66 45 N'toc 

(MNHN) 
Holo* 

14.5 7.1 21.4 69 46 Sibiti 

(SMNH) 

13.9 7.2 21.3 69 48 Lukula 

(MRAC) 
212.583 f 

14.0 6.5 20.1 69 47 Kola 

(MRAC) 
196.340 

14.5 7.1 20.5 72 51 Kola 

(MRAC) 
196.341 

11.6 6.0 16.4 69 51 Kayes 

(MRAC) 
214.004 f 
11.8 6.3 15.1 72 64 Kola 

(MRAC) 
791.389 



Total specimens examined: 7. Sources: MNHN. MRAC, SMNH. 

more than a subspecies of L. mollicella, as Bequaert & 
Clench concluded (1934b:273). This case is reminiscent of the 
conchologically very similar but anatomically contrasting 
Achatina reticulata Pfeiffer, 1845 and A. albopicta E.A. 
Smith, 1878 (Mead, 1950:232). 

Jousseaume's illustration of the holotype is misleading 
because the artist has shown the apex acuminate; actually, 
the first and second whorls are noticeably larger, producing a 
narrowly obtuse apex. 

Apparently neither Leonardo Fea (Germain, 1916) nor 
Captain Vignon (Ancey, 1888) went far enough south in 
Gabon to encounter true L. petitia. 

ACHATININAE 

Bequaertina new genus 

Thin, fragile, anomphalous, medium to large, ovate to ovate- 
elongate shells, 40-80 mm in length. Spire tends to be 
mammillate, apex obtuse. Aperture large, columella long and 
slender, squarely or obliquely truncated. Whorls 6-6V2, rarely 
7; second and third nepionic whorls sculptured; last whorl 
ventricose, ca. 80% of shell length. Sculpture may be vari- 
ously cancellate-granulate, lirate, malleate or nearly smooth. 
Surface of shell lusterless; its abrasion reveals a brilliant inner 
periostracal layer. An occasional specimen may show in the 
periostracum of the lower whorls limited areas of an 
extremely fine decussate micromesh, commonly seen in a 
wide variety of achatinids. 

The generic characters in the soft anatomy are based on 
features that are shared by the two available species - B. 
pintoi (Bourguignat, 1889) and B. graueri (Thiele, 1911). 
Because of the similarity in the basic anatomical pattern in 
these two species, and because, on the basis of shell charac- 
ters, each of the species represents a different dichotomous 
group, it is felt that the following anatomical characters will 
prove to be valid for the genus. 

The most prominent features of the genital system are the 
long free oviduct, the apical vas deferens and the large, 



elongate sacculate spermatheca - all held in close juxtaposi- 
tion by a distinct fascia. In contrast, the penis and penis 
sheath are inconspicuous. The penis sheath enshrouds a short 
basal portion of the long vas deferens. Without exception in 
26 dissected specimens, the penial retractor inserts on or near 
the diaphragm where the latter joins the mantle and the body 
wall of the neck region. At the origin of the penial retractor, 
muscle fibrils pass snugly over the apical penis and then fan 
out into a network that covers the basal vas deferens and the 
inner wall of the penis sheath, except for a limited smooth, 
shiny zone on the approximately upper half of the left side. 
Below this, the fibrils infuse intimately with the tissues of the 
basal penis and penis sheath to create an ill defined section of 
the male conduit that contains the penial atrium. This atrium 
connects the lumen of the penis with the genital atrium. At 
this level, abundant hypertrophied eversion muscle bands 
obscure the genital atrium and its junction with the male and 
female conduits. There is no pilaster or verge. 

The spermathecal duct is much shorter than the spermath- 
eca and often sessile on the vagina. This significantly places 
the spermatheca in a basal position with its usually attenuated 
apex stretching to its connection by fascia to the junction of 
the free oviduct and the apical vas deferens, well below the 
spermoviduct. The hermaphroditic duct has in its midsection 
an enlarged glandular structure of unknown function. The 
ductules to the five hermaphroditic acini are inordinately 
gross and highly convoluted. 

The eggs are covered with a hard calcareous shell and are 
proportionately larger than those of Achatina, i.e. on a par 
with those of Tholachatina (sensu Bequaert, 1950). There is 
no evidence of ovo viviparity. 

The anterior aorta is on the floor of the diaphragm and 
passes ventrally through the diaphragm to the sagittal myo- 
septum in the haemocoele. The second largest vein in the 
lung drains the region near the extreme left mantle and joins 
the primary vein near the apex of the kidney. The large last 
whorl of the shell allows for a highly vascularized left side of 
the lung. The secondary ureter is completely closed. The 
rachidian tooth of the radula is either slender and question- 
ably functional or broad and about half the size of the 
adjacent laterals. The jaw is narrow and broadly arcuate, 
with many slender vertical ribs irregularly distributed. 

Six specimens of B. pellucida (Putzeys, 1898) and one of B. 
marteli (Dautzenberg, 1901), as well as several of Achatina 
craveni, have been found with a single, almost perfectly 
circular hole, 0.6-4.0 mm in diameter, cut usually in the 
dorsal part of the last whorl. These are thought to be caused 
by bird pecks (Meredith, 1983a:25). 

The five species in this genus were place in the genus 
Callistoplepa on the basis of similar shell characters: thin 
shell, large aperture and a tendency to form a mammillate 
spire (Pilsbry 1919, Bequaert & Clench 1934c). But as shown 
in the Key to Subfamilies, a study of the internal anatomies 
revealed major differences. Bequaertina reflects strongest 
phylogenetic affinities to subgenus Achatina (sensu Bequaert, 
1950), particularly with respect to the configuration of the 
basal male conduit and to the fact that the spermatheca is 
attached to the adjacent free oviduct and apical vas deferens 
rather than to the spermoviduct. In Bequaertina, the apical 
penis is free from the apical penis sheath and therefore can 
evert independently, with the sheath following seriatim at 
extroversion, whereas in subgenus Achatina the penis is 
completely enmeshed in a dense network of muscle fibrils and 
connective tissue that requires the penis and the sheath to 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



19 




Figs 23, 24 Callistoplepa barriana (Sowerby, 1890); lectotype Achatina barriana BMNH no. 1889.11.19.2. 25, 26 C. shuttleworthi (Pfeiffer, 

1856); lectotype Achatina shuttleworthi BMNH. Bar scale + ~ 10 mm. 
Figs 27, 28 Leptocala mollicella (Morelet, 1860); lectotype Achatina mollicella BMNH no. 93.2.4.119. 29, 30 L. petitia (Jousseaume, 1884); 

holotype Petitia petitia MNHN. 31, 32 Bequertina pellucida (Putzeys, 1898); lectotype Ganomidos pellucidus MRAC no. 5135. 33, 34 B. 

pellucida; paralectotype G. pellucidus (unicolorous) MRAC no. 5136. 35, 36 B. pellucida; lectotype Serpaea foai Germain, 1905 MNHN. 

37, 38 B. marteli (Dautzenberg, 1901); lectotype Achatina marteli IRSN. 39, 40 B. marteli; lectotype A. marteli pallescens (Dautzenberg, 

1901) IRSN. 41, 42 B. pintoi (Bourguignat, 1889); holotype, Serpaea pintoi MNHN. Bar scale = ~ 10 mm. 
Figs 43, 44 B. pintoi; lectotype Achatina fragilis Smith, 1899 BMNH no. 97.12.31.9. 45, 46. B. pintoi; holotype, Callistoplepa thielei, 

Bequaert & Clench, 1934c ZMB no. 53177. 47, 48 B. pintoi; BMNH no. 1885.5.25.47. 49, 50 B. pintoi; BMNH MacAndrew (1563). 

51 B. pintoi; BMNH no. 1907.7.25.3. 52, 53 B. fraterculus (Dupuis & Putzeys, 1900); lectotype Ganomidos fraterculus MRAC no. 5140. 

54, 55 B. graueri (Thiele, 1911); lectotype Achatina graueri ZMB no. 101935. 56, 57 B. graueri; lectotype Callistoplepa babaulti, 

Germain, 1936 MNHN. Bar scale = ~ 10 mm. 



20 



A.R. MEAD 







NEW SUBFAMILY AND GENUS ACHATINIDAE 



21 



£*C^^rV ^F ^ 


^^ >«^^r ^B 


E .x ■ 




M 1 


^^^^Hr fl^^^^H ^^BR^* 

V t ^H 1 


jj^H), V Bar 

11 J 


^^^HF J El 


■^^^fl ^jfl 



22 



A.R. MEAD 



evert together as a unit, forming a quite different intromittent 
organ. The very shell characters that seemed to link these five 
species with Callistoplepa now are seen conchologically to 
distinguish the more primitive genus Bequaertina from Acha- 
tina s.s. 

On the basis of somewhat overlapping shell characters, 
available distributional records, and the limited anatomical 
evidence, the species of Bequaertina break into two groups: 
(1) the malleate, mammillate B. fraterculus (Dupuis & 
Putzeys, 1900) and B. graueri, and (2) the cancellate B. 
pellucida, B. marteli and B. pintoi. B. pellucida of southeast 
Zaire, close to what is believed to be its ancestral home, is 
plesiomorphic within the group and stands between an ances- 
tral achatinid stock of the Zaire Basin and Achatina s.s., 
which today is largely restricted to that basin. A branch of the 
ancestral stock moved north and northeast to give rise to the 
apomorphic B. fraterculus and B. graueri. A more conserva- 
tive second branch moved east to give rise to B. pellucida and 

B. marteli. This second branch continued further east and 
then into a strong north-south axis to give rise to the closely 
related B. pintoi. The known distribution of the genus (Fig. 
16) embraces a vast area of the Rift Valley - Lake Region and 
the Lualaba branch of the Zaire River in central, eastern and 
southeastern Africa. Greater field collecting will probably 
extend the limited distributions of B. fraterculus and B. 
marteli. 

Pilsbry & Cockerell (1933), on the erroneous assumption 
that Achatina graueri 'represented an intrusion of a South 
African type into the Central African region', initially 
decided to place it in the genus Cochlitoma. They softened 
their stand on the advice of M. Connolly and designated it 
'Achatina (Cochlitoma) graueri'. However, this species can- 
not possibly be considered congeneric with Cochlitoma zebra 
(Bruguiere, 1789), which Pilsbry (1904:xiii, 78) selected as 
the type species of the genus Cochlitoma, because Mead 
(1992) shows Achatina zebra anatomically belongs to subge- 
nus Tholachatina of Archachatina. Since the present group of 
five species is not congeneric with the species in either 
Callistoplepa or Achatina s.s., and since this group also is not 
congeneric with Achatina hortensiae Morelet, 1866, which 
Pilsbry (1904:21) selected as the 'type' of Serpaea, there is no 
other available genus-group name. Ganomidos cannot be 
considered because it is a junior subjective synonym of 
Callistoplepa. For these reasons, the generic name Bequaer- 
tina is proposed. It is named in honour of the late Dr Joseph 

C. Bequaert, Agassiz Professor of Zoology at Harvard Uni- 
versity, who will remain the classical authority not only in the 
Achatinidae, but also in several families of insects and 
arachnids that he mastered in his long lifetime. Because B. 
graueri is the largest and most conspicuous of the five species, 
and because it has departed farthest from what is believed to 
be the ancestral stock, it is here selected as type species of the 
genus. 

Early in the present study, it became obvious that this 
group of five species anatomically was not congeneric with 
Callistoplepa. This information was shared with colleagues 
who considerately referred to this new genus in general terms 
(van Bruggen, 1978:912, 921, 1988:10; van Bruggen & 
Meredith, 1984:161). Also, the present author made refer- 
ence to this new genus in an earlier manuscript as 'Callis- 
toplepa s.i: (Mead, 1992). 



Key to Species 

1 Last whorl distinctly granulate above the periphery, or at least 
in a limited subsutural zone; growth wrinkles conspicuous to 
dominant 2 

Last whorl faintly granulate, malleate, lirate or smooth except 
for modest irregular growth wrinkles 4 

2 Apex broadly obtuse; 6 whorls = > 50 mm; yellowish, ochra- 
ceous or olivaceous; gross granulate sculpture; first nepionic 
whorl 2-3 mm in diameter; second whorl expanding broadly; 
sculpture of second whorl coarse and either distinctly granular 
or depressed and poorly defined; transverse measurement at 
junction of third and fourth whorls is 2'/2-4 mm; outer lip 
increasingly arcuate basally; growth wrinkles bold or moder- 
ately heavy 3 

Apex subacute to narrowly obtuse; 6 whorls = ~ 40 mm, 7 
whorls = 53-60 mm; translucent dull fulvous to dull olivaceous- 
brown; moderately coarse to fine granulate sculpture; first 
nepionic whorl 2 mm in diameter; second whorl tends to be 
slightly constricted, expanding limitedly; sculpture of second 
whorl finely engraved, delicate; transverse measurement at 
junction of third and fourth whorls is 2-2'/:? mm; outer lip evenly 
arcuate; growth wrinkles thin, of modest calibre. Southeast 
Zaire, northeast Zambia and west central Tanzania . pellucida 

3 Last whorl large, rarely strikingly so; ground colour intense 
olivaceous-yellow to subdued olivaceous; prominent closely 
aligned somewhat irregular costate transverse ridges embrace 
the gross elongate granules with bold vertical emphasis, domi- 
nating the spiral lines; strongly contrasting zigzag castaneous 
flammules usually present, pale unicolorous forms uncommon; 
first nepionic whorl 2V2-3 mm in diameter; sculpture of second 
whorl coarse, granular, elevated, tightly packed; transverse 
measurement at junction of third and fourth whorls is 3-4 mm; 
third whorl deeply and grossly granulate. Middle west and east 
shores of Lake Tanganyika, Zaire and Tanzania marteli 

- Last whorl large, often very large to ventricose; ground colour 
dark olivaceous to pale olivaceous yellow; coarse granulate 
sculpture above periphery, reduced or absent below periphery 
(varies within a single whorl); transverse ridges moderate, 
slender, fairly uniform, in balance with the spiral lines, con- 
spicuous below periphery but obscured by granulate sculpture 
above periphery; usually unicolorous, but narrow fairly straight 
light castaneous stripes may be present; first nepionic whorl 
2-2'/2 mm in diameter; sculpture of second whorl coarse, but 
superficial, vaguely and irregularly impressed, patchy, poorly 
defined, often worn smooth; transverse measurement at junc- 
tion of third and fourth whorls is 2V2-3 mm; third whorl 
delicately to moderately granulate. East Africa, almost reaching 
the Limpopo River in the south (4-20° S, 27-39° E) pintoi 

4 Apex of shell obtuse and noticeably mammillate; shell conspicu- 
ously to obscurely malleate; opaque or dark and translucent, 
uniformly or somewhat variably brown or yellow-brown, band- 
ing limited and irregular; coarse growth wrinkles or extremely 
fine lirae dominate the sculpture, 6 whorls = > 43 mm 5 

Apex of shell subacute to narrowly obtuse, somewhat elevated 
but not mammillate; shell not malleate; translucent dull fulvous 
to dull olivaceous-brown, usually with moderately broad casta- 
neous flames and stripes irregularly distributed, but may be pale 
unicolorous; very fine granulate-cancellate sculpture dominates; 
6 whorls = < 43 mm. Southeast Zaire, northeast Zambia and 
west central Tanzania pellucida 

5 Shell large (6'A whorls = 60-80 mm), thin but substantial, 
essentially opaque; usually conspicuously malleate; not cari- 
nate; growth wrinkles prominent, rather regular; lirae of fifth 
whorl distinctly transacted by spiral striae; unicolorous or 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



23 



Table 5 Locality records - Bequaertina. Numbers in the first 
column correspond to the locality numbers in Figure 16. Sources 
of specimen information are shown in the last column. 



1. Lake Tanganyika, 1800-2000m 4° 30' S, 29° 00' E 

2. Kiambi 7° 20' S, 28° 01' E 



3. Sampwe (non 'Sangue') 



9° 20' S, 27° 26' E 



4. Ibahi, Ugogo ( = Ougogo) Riv. 5° 04' S, 34° 04' E 

5. Mbwe( = Mbwego) 5° 21' S, 38° 58' E 

6. Mamboya ( = Mamboa) 6° 16' S, 37° 06' E 

7. Morogoro 6° 50' S, 37° 45' E 

8. Ngerengere, Oukani, Kingoni 7° 03' S, 38° 31' E 



9. Ufipa ( = Sumbawanga) 

10. Rukwa Lk. 

11. Mbaya, 1700m 

12. Utengule 



13. Misuku Hills, Mughoma, 

1500m 

14. Deep Bay ( = Chilumba, 

= Hengwa) 

15. Nyika Plateau, 6000-7000 ft. 10° 48' S, 33° 48' E 



NHMW* 
Dautz. & 
Germ., 
1914 

Dautz. & 
Germ., 
1914 
Ancey, 
1902 
Ancey, 
1902 
BMNH 
BMNH 
Bourg., 
1889 
Ancey, 
1902 
BMNH f 
BMNH, 
LNK 
BMNH, 
MCZ, 
ZMB, 
SMF 
9° 40' S, 33° 33' E RMNH + 

10° 27' S, 34° 16' E BMNH 



8°00'S, 31°30'E 

8° 00' S, 32° 25' E 
8° 45' S, 33° 27' E 

8° 54' S, 33° 20' E 



16. Nkota-Kota 

17. Nchisi ( = Ntchisi) Mt. 

18. Chinyama 

19. Zomba, Shirwa Lk., Mpita 



12° 55' S, 34° 18' E 
13° 20' S, 34° 05' E 
13° 43' S, 33° 43' E 
15° 23' S, 35° 23' E 



20. ChiradzuluMt., Lisau 15° 41' S, 35° 09' E 

21. Nyambadwe Hill 15° 48' S, 35° 15' E 

22. SocheMt. 15° 51' S, 35° 01' E 

23. Cheri Bridge, Upper Lauangwal3° 35' S, 31° 30' E 

24. Broken Hill ( = Kabwe) 14° 27' S, 28° 27' E 

25. Kafue Riv., Mumbwa 15° 56' S, 28° 55' E 

26. Pemba 16° 40' S, 27° 25' E 

27. Mazoe Valley 16° 32' S, 33° 25' E 

28. Salisbury 17° 50' S, 31° 03' E 

29. Vumba, Zonwi Bridge, 2500 ft. 19° 07' S, 33° 05' E 

30. Bulawayo 20° 09' S, 28° 35' E 

31. Chirinda, Selinda Mt., 4000 ft. 20° 26' S, 32° 42' E 

32. Macequece, Vila de Manica 18° 56' S, 32° 53' E 



33. Nsendwe, Maniema 

34. Uvira 

35. Mpala(=Pala) 

36. Mweru ( = Moero) Lk. 

37. Dilolo 

38. Rumonge 

39. Kapuri ( = Piani Kapuri) 



40. Luaye 



2° 57' S, 25° 56' E 

3° 24' S, 29° 08' E 
6°45'S,29°31'E 
9° 00' S, 28° 45' E 

10° 42' S, 22° 20' E 
3° 11' S, 29° 08' E 
3° 34' S, 26° 53' E 



4° 42' S, 27° 23' E 



BMNH, 

IRSN, 

MCZet 

al. 

BMNH 

HM' 

HM* 

BMNH, 

IRSN, 

RMNH 

HM* 

NG f 

NG f 

MCZ 

NMW 

Beq. & 

CI., 1934c 

SAM 

NMW 

NM 

NM* 

RMS, 

SAM 

BMNH, 

MCZ 

BMNH, 

NMW, 

SAM 

BMNH, 

MRAC 

ZMB 

IRSN 

BMNH, 

IRSN 

SMF 

MRAC 

BMNH, 

IRSN, 

MRAC et 

al. 

MRAC 



variably transversely striated with yellow-brown to dark brown; 
nepionic whorls densely granulate. Lake Kivu district of Zaire, 
Rwanda, Uganda graueri 

- Shell small (6 1 /-* whorls = ~ 50 mm), extremely thin, fragile, 
translucent; malleations very shallow, often sparse; subcarinate 
at periphery, producing a bend in the arc of fine prosocline 
lirae; lirae of fifth whorl not transacted; ground colour dull dark 
brown-olive with irregular castaneous brush marks that are 
closely highlighted adaperturally with buff; nepionic whorls 
faintly granulate. Lualaba River, Zaire fraterculus 

Bequaertina pellucida (Putzeys, 1898) 
Figs. 31-36 

Ganomidos pellucidus 

Putzeys, 1898:84, text fig. 20, 21. 
Callistoplepa pellucida 

Pilsbry, 1905:128, pi. 43, fig. 3, 4; Germain, 1909:90; 

Pilsbry, 1919:81; Bequaert & Clench, 1934c:114; Haas, 

1936:13. 
Serpaea foai 

Germain, 1905:255; 1908:631. 
Achatina foai 

Verdcourt, 1966:111; 1983:219. 
Callistopepla pellucida 

Oliver, 1983:9. 

Shell. Shell ovate-achatiniform to elongate-ovate, 
extremely thin, very fragile, translucent. Whorls 6-7, rarely 
l x h\ a conspicuous demarcation at or near the end of the third 
whorl sets off the nepionic whorls. Spire conic, with a 
narrowly obtuse apex that is slightly elevated; occasionally 
the second nepionic whorl is somewhat constricted, produc- 
ing a submammillate profile. Whorls slightly convex, expand- 
ing and descending proportionately. Sutures moderately deep 

Table 5 continued 



41. 


Kabambare 


4° 42' S, 27° 43' E 


ZMB 


42. 


Lukuga Riv. mouth 


5° 55' S, 29° 12' E 


MCZ 


43. 


Gandajika 


6° 45' S, 23° 57' E 


MRAC 


44. 


Pweto 


8° 26' S, 28° 55' E 


IRSN 


45. 


Kamina 


8° 44' S,25°00' E 


MRAC 


46. 


Abercorn ( = Mbala) 


8°50'S, 31°22'E 


MCZ 


47. 


Kungwe, Sitete ( = Nkungwe) 


6° 07' S, 29° 48' E 


Verdcourt, 
1966 


48. 


Beni 


0°30' N,29°28' E 


IRSN, 
MRAC 


49. 


Kitembo 


2° 53' S, 27° 37' E 


MNHN 


50. 


Lobengera Mission 


2° 03' S, 29° 25' E 


MRAC 


51. 


Ibanda 


0° 08' S, 30° 30' E 


MRAC 


52. 


Loashi Valley 


1° 14' S, 28° 45' E 


MRAC 


53. 


Burungu, Ruasa 


1° 20' S, 29° 02' E 


ANSP 


54. 


Nyabukere 


1° 29' S, 28° 33' E 


MRAC 


55. 


Kirotche, 1250 m 


1° 37' S, 29° 02' E 


MRAC 


56. 


Lwiro Riv. 


2° 00' S, 28° 52' E 


AMNH, 

FMNH 


57. 


Idjwi ( = Kwidschwi, Kwidjwi) 


2° 09' S, 29° 04' E 


ZMB, 
MRAC, 
UMMZ et 
al. 


58. 


Katana 


2° 13' S, 28° 50' E 


MRAC 


59. 


Tshibinda 


2° 20' S, 28° 45' E 


ANSP, 
MRAC 


60. 


Bukavu 


2° 30' S, 28° 52' E 


MNHN, 
NMB 



specimens dissected in the present study. 



24 

and irregular. Last whorl large, 80% of shell length; range for 
5-7V2 whorls, 76-84% (n = 52). Aperture oval, faint milky 
wash within. Columella concolorous, slender, moderately 
long straight or slightly arcuate and rectangularly to very 
obliquely truncate. Outer lip extremely thin, evenly arcuate, 
receding at base in profile. Parietal callus scarcely detectable. 

The first two to three whorls are light horn colour. Pale, 
obscure, castaneous streaks begin to appear in the third or 
fourth whorl; these characteristically are broader at the 
suture below, becoming increasingly darker, larger and more 
irregular on the last whorl. These streaks may be variously 
vertical, diagonal, angulate, flammulate, interrupted, or 
reduced to spots and blotches. Ground colour is dull buff to 
dull olivaceous-fulvous. Of 56 specimens checked precisely 
for colour, 63% have a definite pattern, 7% are nearly 
unicolorous, and 30% are unicolorous. There was no correla- 
tion between colour pattern and locality. 

The first whorl is essentially without sculpture. Minute, 
faintly engraved crescentic granulations usually appear early 
in the second whorl; these are formed by nearly equidistant 
spiral lines and irregular, scalloped transverse lines. The 
latter become straighter and compressed in the third whorl, 
producing narrow elongate granulations and irregularly 
appearing prosocline growth wrinkles that are crenulate at 
the suture. The sculpture becomes more disperse in the 
fourth and fifth whorls, producing a dominant, fairly uni- 
form, subquadrate, often welt-like, cancellate-granulate 
sculpture, which usually fades quickly at the periphery. In the 
sixth to seventh whorls, this sculpture becomes more and 
more subdued and diffuse until the increasingly prominent, 
yet modest, growth wrinkles dominate both above and below 
the periphery. In the largest specimens of seven whorls the 
cancellate-granulate sculpture may feebly or strongly return 
both above and below the periphery. The dull, extremely 
thin, tenaceous outer periostracal layer wears off in very 
limited areas, highlighting the sculpture with the exposed 
glossy inner periostracal layer. 

Soft anatomy. No known alcohol preserved specimens. 

Type material. As nearly as can be determined, Putzeys 
had 14 syntypes of his Ganomidos pellucidus, for which he 
gave a range of shell dimensions (1898). The specimens were 
collected by P. Dupuis. Putzeys retained a select series of 7 
syntypes in his own collection (MRAC no. 5132-5138). He 
did not designate types, but selected the largest specimen 
(no. 5132) for an abaperatural view and a small specimen with 
slender flames (no. 5133) for an aperatural view in his line 
drawing illustrations. Regrettably, the larger specimen had 
been rather badly damaged and mended in nature, and the 
smaller specimen was excessively small. The second largest 
syntype in his series (no. 5136) is a unicolorous specimen that 
is representative of only about a quarter of the known 
specimens (Figs. 33, 34). Hence, the flamed, third largest 
syntype in his series (no. 5 135) is here selected the lectotype 
(Figs. 31, 32; Table 6), with the other syntypes becoming 
paralectotypes (BMNH lno. 1904.5. 18.68, IRSN 5, MRAC 6, 
NHMW 1). 

On the basis of two specimens collected by Edouard Foa 
during his 1897-98 expedition to the Lake District of Africa, 
Germain described (1905) and figured (1908) the junior 
subjective synonym Serpaea foai from 'Tanganika est', later 
corrected to 'les bords du Lac Tanganyika'. Bequaert (1950) 
placed Serpaea in the synonymy of Achatina, but apparently 



A.R. MEAD 
Table 6 B. pellucida - Representative shells measurements. 







Greatest Aperture 


Last 


/o 




Whorls 


Length 


Width 


Length Width whorl LW/L % W/L 


7 


60.4 


31.4 


32.7 


19.5 


45.9 


76 


52 Mweru 
(BMNH) 


7Vz 


58.6 


32.8 


36.0 


16.6 


46.0 


79 


56 Kamina 
(MRAC) 
581.196 


7 


54.5 


29.0 


30.5 


17.0 


42.5 


78 


53 Mweru 
(BMNH) 
1907.11. 
11.5 


7 


53.3 


30.2 


31.2 


17.8 


42.3 


79 


57 Mpala 
(IRSN) 


6V2 


48.0 


28.0 


29.7 


15.8 


38.0 


79 


58 Piani 
Kapuri 
(MRAC) 
5132 PLec 
G.p. 


6V2 


46.9 


29.6 


29.4 


16.5 


37.5 


80 


63 Tanganyika 
(MNHN) 
Lect 5. 
foai * 


6V4 


45.8 


28.0 


28.0 


16.7 


37.0 


82 


61 Piani 
Kapuri 
(IRSN) 
PLec G.p. 


6V2 


45.0 


28.2 


26.8 


15.5 


35.5 


79 


63 Piani 
Kapuri 
(MRAC) 
5136 PLec 
G.p.* 


6V4 


43.2 


24.6 


25.5 


14.1 


33.9 


78 


57 Piani 
Kapuri 
(MRAC) 
5135 Lect 
G.p.* 


6V< 


40.4 


22.6 


23.0 


13.6 


31.7 


78 


56 Piani 
Kapuri 
(MRAC) 
5133 PLec 
G.p. 


6 


39.0 


23.3 


24.4 


14.1 


32.0 


82 


60 Tanganyika 
(MNHN) 
PLec S. 
foai 


5% 


35.5 


20.0 


20.9 


12.0 


28.4 


80 


56 PLec G.p. 
(BMNH) 
1904.5. 
18.68 



Total specimens examined: 62. Sources: BMNH, IRSN. MCZ, MNHN, 
MRAC, NMW, USNM, ZMB. 



overlooked Germain's species. Only Verdcourt (1966) has 
acknowledged the existence of this species, and then only as 
an East African species unknown to him. A study of the two 
syntypes in Paris (MNHN) confirmed the fact that they are 
indeed Putzeys' species. His larger, sharply photographed 
'seul adulte' specimen (Figs. 35, 36) is here selected as the 
lectotype of Germain's Serpaea foai (Table 6). 

Deshayes (1824-37, 1864) described and illustrated a small 
fossil snail Agathina pellucida (= Achatina pellucida) from the 
Paris basin. Lamarck (1838:313) also refers to this species. 
This very acuminate, slender specimen is possibly a subulinid. 
It does not enter into homonymy with Putzeys' G. pellucidus 
because the latter was never included in the genus Achatina. 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



25 



Type locality. Forest of Piani Kapuri, Maniema 
(=Manyema), Zaire 3° 34' S, 26° 53' E. 

Distribution. Next to B. pintoi, this is the most wide 
spread species in the genus (Fig. 16). The known specific 
localities delineate essentially the southeastern quarter of 
Zaire, with Nsendwe, near Kindu-Port-Empain, Maniema 
region in the northwest; Uvira, Kivu region in the northeast; 
Mpala, Tanganyika region in the east; Lake Mwero 
( = Moero), Kantanga region in the southeast; and Dilolo, 
Lualaba region in the southwest. The only records outside 
Zaire are (1) in Abercorn (= Mbala) at the southern tip of 
Lake Tanganyika, Zambia, and (2) on the Tanzanian east 
shores of this lake, based on Germain's synonym Serpaea foai 
(1905, 1908). Meredith (1983b) and N. Gray (correspon- 
dence) failed to find it during extensive collecting in Malawi. 
The largest and finest specimens extant were collected in the 
Lake Mwero region (BMNH) and Kamina (MRAC). 

Remarks. This plesiomorphic wide spread species is most 
closely related to B. marteli. Specimens have been found in 
mixed lots along with B. marteli and Achatina craveni. The 
juvenile specimens of all three species are easily confused. 
Further, the full grown specimens are quite variable in shape, 
colour, sculpture and pattern, with the not uncommon atypi- 
cal forms of each species contributing to the difficulty of 
identification. The young specimen that Grauer collected in 
the virgin forest 50 km east of Kasongo, Zaire, and identified 
as Achatina fulminatrix von Martens, 1895 by Thiele 
(1911:205) was examined in Berlin (ZMB) and found to be B. 
pellucida. Extensive series of this species are in Bruxelles 
(IRSN) and Tervuren (MRAC). 

tit quae it in a marteli (Dautzenberg, 1901) 
Figs. 37-40 

Achatina marteli 

Dautzenberg, 1901:3. 
Ganomidos marteli 

Dautzenberg, 1901, pi. 1, fig. 1. 
Achatina marteli pallescens 

Dautzenberg, 1901:3. 
Ganomidos marteli pallescens 

Dautzenberg, 1901, pi. 1, fig. 2. 
Callistoplepa marteli 

Pilsbry, 1905:129, pi. 47, fig. 21 (ex Dautzenberg); Ger- 
main, 1909:90; Pilsbry, 1919:81; Bequaert & Clench, 

1934c:114. 
Callistoplepa marteli var. pallescens 

Pilsbry, 1905:129, pi. 47, fig. 22 (ex Dautzenberg); 

Bequaert & Clench, 1934c: 114. 
'Achatina sp. near tavaresiana 

Verdcourt, 1966:106, fig. 12; 1988:219. 
Callistopepla marteli 

Germain, 1936:151; Oliver, 1983:9. 

Shell. Shell ovate-achatiniform, opaque, thin but not frag- 
ile. Whorls 6-6V4, rarely 6V2. Spire moderately broad, conic; 
apex obtuse; only one out of 69 specimens examined had a 
mammillate apex. Upper whorls only slightly convex, 
descending proportionately but expanding somewhat more 
rapidly. Sutures fine and regular in nepionic whorls, shallow 
to moderately deep and irregular in the following whorls. 
Last whorl large and more convex, 80% of shell length; range 
for 4V2-6V2 whorls, 77-84% (n = 69). Aperture inverted 



auriform to ovate-elongate; pale blue-white within; surface 
pattern and flames show through. Columella straight or 
weakly arcuate, somewhat slender, concolorous but with a 
thin calcareous film; usually moderately obliquely truncated. 
Outer lip thin, extending basally only a slight way below the 
truncation; its arc is characteristically greatest below midway 
in the mature specimens. Parietal callus thin but apparent 
even in the smaller specimens. 

The nepionic whorls (first 2V2) are unicolorous pale buff- 
white. This changes imperceptibly to a uniform dull ground 
colour that varies in specimens from a rather intense oliva- 
ceous yellow to subdued olivaceous. In most specimens, 
faint, very diffuse light castaneous blotches appear in the 
fourth whorl. At first these are vertical, evenly spaced and 
broader at their base; but they soon become fragmented 
apically, darker, and strikingly distorted into diagonal even 
spiral, irregular streaks, bands and flames that are approxi- 
mately as wide as the ground colour space between them. In 
the present study of 69 specimens, 72% are flammate, 13% 
are vaguely flammate but only on the last whorl, and 15% are 
without flames, i.e. 'pallescent'. In some of the latter, e.g. the 
lectotype of Achatina marteli pallescens, lines of arrested 
growth are highlighted with thin bands of dark brown. 

A delicate beaded or slightly semilunar sculpture starts in 
the second quarter of the first whorl and quickly assumes in 
the early second whorl the diagnostic sculpture of strikingly 
coarse, elevated, round or crescentic, discreet but tightly 
packed beads that are neatly aligned in 5-7 spiral rows. This 
pattern persists almost uniformly throughout the second 
whorl. In the mid-third whorl, the transverse rows become 
greatly compressed, producing growth wrinkles and convert- 
ing the beads into transverse welts 2-3 times as long as wide. 
This doubtless marks the first postemergent growth. Adaper- 
tural to this, the growth wrinkles become prosocline, the 
sculpture gradually becomes less compressed, the spiral striae 
become more numerous and deeper, and the individual welts 
become larger, more variable in size, more rectangular, and 
often cleft. The remarkably evenly and closely spaced coarse 
growth wrinkles embrace and intensify the prosocline rows of 
welts, producing the characteristic prominent ribbed sculp- 
ture of this species. Apically, the ribs may bifurcate and form 
crenulations. Below the periphery, the welts rather abruptly 
reduce to one-quarter their calibre, or are absent, leaving 
prominently the growth wrinkles. An extremely fine decus- 
sate micromesh of the periostracum appears on the last whorl 
of some specimens. It is more noticeable on the shiny inner 
layer of the periostracum where the latter is exposed through 
wear or injury. It is apparent that the micromesh is formed at 
the time that the inner periostracal layer is laid down and that 
it is largely obscured by the preformed, smoother outer 
periostracal layer. It is likely that the micromesh assists 
structurally in bonding the two periostracal layers. 

Soft anatomy. No known alcohol preserved specimens. 

Type material. In his description of this species and its 
synonymous unicolorous 'variety pallescens\ Dautzenberg 
(1901) announced that he was dedicating them to Colonel 
Martel and that specimens had been collected by R.P. 
Guilleme 'en nombreux exemplaires' in the region of Lake 
Tanganyika. He did not specifically designate types and 
paratypes, although he selected a fine flamed specimen and 
an equally fine unicolorous specimen that were photo- 
graphed, both in apertural view only, as representative of the 



26 



A.R. MEAD 



two proposed taxa. These are in the type collection in 
Bruxelles (IRSN) and are here selected as lectotypes of 
Dautzenberg's Achatina marteli and A.m. pallescens, respec- 
tively (Figs. 37-40; Table 7). As he pointed out in a footnote 
in the original descriptions, the pronounced flame pattern of 
his figure 1 unfortunately did not reproduce well. Pilsbry's 
copies (1905) therefore reflected this deficiency. In this 
species, neither the lack of colour pattern nor the greater 
degree of ventricosity is taxonomically valid for establishing a 
trinomen. 

In the IRSN collection there are several mixed lots totalling 
48 mostly juvenile, damaged or weathered specimens. All 
these specimens were very carefully examined in the present 
study and were found to be a mixture of the flamed and 
unicolorous forms of this species and, in addition, juveniles of 
Bequaertina pellucida and Achatina craveni. These cannot 
reasonably be considered to have been a part of Dautzen- 
berg's type series. Dautzenberg, however, did distribute his 
specimens widely. Those bearing the type locality and R.P. 
Guilleme as the collector are here selected as paralectotypes. 
The known distribution of these flamed/unicolorous speci- 
mens are BMNH 1/0, NMW 1/1, IRSN 8/5, MCZ 2/1, MRAC 
7/1, NMB 2/1, MNHN 6/1, NHMW 1/0. 

Table 7 B. marteli - Representative shells measurements. 



Greatest Aperture Last % 

Whorls Length Width Length Width whorl LW/L % W/L 



6V2 68.3 34.0 42.3 20.8 55.4 81 50 Mpala 

(IRSN) 
PLec 

6V4 67.2 38.6 39.1 22.2 54.4 81 57 Mpala 

(IRSN) 
Lect 
A.m. p.* 

6 63.7 33.2 37.7 19.6 51.5 81 52 Mpala 

(IRSN) 
Lect 
A.m.* 

6'/4 62.9 32.3 36.7 18.2 49.8 79 51 Mpala 

(MCZ) 
97929 
PLec 

6 60.8 34.8 38.0 19.4 51.3 84 57 Mpala 

(MNHN) 
PLec 

5% 58.4 32.3 37.3 18.5 49.0 84 55 Mpala 

(MRAC) 
5129 PLec 

654.032.233.817.043.78160 

54.0 32.2 33.8 17.0 43.7 81 60 Mpala 

(MNHN) 
PLec 

5% 48.4 26.9 32.1 15.0 39.8 82 55 Mpala 

(BMNH) 
PLec 
1937.12. 
30.1934 

5>/2 48.4 27.9 32.0 16.7 40.5 84 58 Mpala 

(MRAC) 
5125 PLec 

6 48.4 27.3 31.0 15.5 39.6 82 56 Mpala 

(MRAC) 
5131 PLec 



Total specimens examined: 69. Sources: BMNH, IRSN, MCZ, MNHN, 
MRAC, NHMW, NMB, NMW. 



Type locality. Mpala (= Pala) 6° 45' S, 29° 31' E, region of 
Lake Tanganyika, Zaire (cf van Burggen, 1988:9). 

Distribution. All specimens examined are from the type 
locality. Verdcourt kindly sent the author a photograph of a 
specimen, which earlier had been identified as Achatina 
tavaresiana (Pain & Verdcourt, 1962; Verdcourt, 1966). This 
specimen is clearly B. marteli and establishes this species in 
Tanzania on the eastern shore of Lake Tanganyika in the 
Mahari Peninsula at Nkungwe (= Kungwe) 6° 07' S, 29° 48' 
E. It was collected in 'litter in thick scrub at head of stream, 
altitude 4500 ft'. With the many locality records known for B. 
pellucida, including Mpala, it is strange that the distribution 
of the present species is, to date, so contrastingly limited. 

Remarks. This is the least fragile and the most boldly 
sculptured species in the genus. Phylogenetically, it appears 
to stand between B. pellucida and B. pintoi. Mixed lots of B. 
marteli, B. pellucida, and Achatina craveni suggest that these 
species are sympatric. The juveniles in particular are confus- 
able. By far the largest series of this species is to be found in 
Bruxelles (IRSN). 

Bequaertina pintoi (Bourguignat, 1889) 
Figs. 41-51 

Serpaea pintoi 

Bourguignat, 1889:86, pi. 4. fig. 4. 
Achatina fragilis 

Smith, 1899:591, pi. 35, figs. 3,4 (non Achatina fragilis 

Deshayes, 1864); Ancey, 1902:278. text fig. 6; Pilsbry, 

1904:63, pi. 9, figs. 25, 26 (ex Smith); Dautzenberg & 

Germain, 1914:26. 
Achatina . . . . sp. nov?' 

Ancey, 1902:277, text fig. 4. 
Achatina pintoi 

Pilsbry, 1904:63, pi. 41, fig. 8 (ex Bourguignat); Bequaert, 

1950:11; Verdcourt, 1966:111, 1983:219. 
Achatina nyikaensis 

Pilsbry, 1909:113, 1919:79; Connolly, 1925:168, 1939:321; 

Germain, 1935:9; van Bruggen, 1965:81, 1988:10; van 

Bruggen & Meredith, 1984:161. 
Callistoplepa nyikaensis 

Bequaert & Clench, 1934c:115, 116. 
Callistoplepa thielei 

Bequaert & Clench, 1934c: 115, pi. 2, figs. 8-10, 12. 
Callistopepla nyikaensis 

Verdcourt, 1966:111, 1983:219; Meredith, 1983a:29, fig. 

10, 1983b:247. 

Shell. Shell thin, fragile, highly variable in shape, usually 
elongate-ovate or ovate-subsuccineiform, but may be ovate, 
globose-ovate or slender conic-ovate. Whorls 6— 6'/2, rarely 
slightly larger. Apex obtuse; vaguely mammillate in some 
specimens. Spire usually inscribes a short broad based tri- 
angle that appears to be nearly equilateral, 27-36% of shell 
length. Less commonly, the spire is more produced, with the 
sides of the connate triangle appearing longer than the width 
of the base, 37-43% of shell length; such specimens may or 
may not have a more slender last whorl. Sutures fine, distinct, 
deeply impressed, quite regular, but may be faintly crenulate 
in the last part of the sixth whorl. Whorls slightly, moder- 
ately, or distinctly convex, usually expanding rapidly to form 
a large last whorl, 84% of shell length; range for 3'/2-6 3 /4 
whorls, 78-87% (n = 87). Aperture ovate-acuminate; charac- 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



27 



teristically widest slightly below middle; translucent pale 
milky opalescent within; fine, closely aligned internal riblets 
mirror the external sculpture. Columella concolorous or 
white; long, slender, feebly arcuate or nearly straight; trun- 
cated obliquely or abruptly. Outer lip extremely thin, fragile, 
somewhat receding, arcuately skewed and evenly rounded 
toward the base. Parietal callus not apparent in young or 
fresh specimens; thinly calcareous white in others. 

Nearly 90% of the specimens examined in the present 
study (77/87) were essentially unicolorous in a colour gradient 
from pale olivaceous yellow to olivaceous brown to deep 
olivaceous green. In individual specimens, the colour tends to 
be quite uniform, except for darker bands where there was 
cessation of growth. Because of their very thin two periostra- 
cal layers, the apical whorls, which are pale straw colour, 
soon become calcareous white with wear and exposure. Ten 
of the specimens examined had on their lower whorls, very 
faint, narrow light castaneous irregular, sometimes inter- 
rupted, transverse stripes that were one-half to one-third the 
width of the ground colour between them. In some speci- 
mens, only a fraction of a whorl was involved. 

Extremely fine spiral engraved lines appear at the end of 
the first whorl. Transverse lines appear in the second whorl, 
giving rise to depressed beads that become more conspicuous 
and more abundant until near the middle of the third whorl. 
At that point, which marks the end of the nepionic whorls, 
the beads become compressed into very narrow transverse 
ridges. In the fourth whorl, this compression is relieved and a 
fairly even granulose-cancellate sculpture emerges. In the 
fifth and sixth whorls, the granulae swell to become welts that 
occasionally anastomose along the transverse growth 
wrinkles, with the shallow spiral lines remaining strongly in 
evidence (cf Bequaert & Clench 1934c, fig. 12). Usually, this 
sculpture diminishes rapidly in caliber at the periphery, with 
essentially only the growth wrinkles continuing into the 
otherwise smooth surface. In other specimens, the sculpture 
may continue strongly below the periphery, but at a reduced 
calibre. In still others, subdued patches of this sculpture 
appear irregularly below the periphery, and perhaps contrast- 
ingly so with resumed growth after diapause. Particularly in 
the parietal area, the dull, tenaceous microscopically granular 
outer periostracal layer may wear off, exposing the smooth, 
shiny inner periostracal layer. Only rarely has a periostracal 
decussate micromesh been observed in this species, and then 
only spottedly below the periphery in the inner periostracal 
layer of the sixth whorl, e.g. in the holotype of Callistoplepa 
thielei Bequaert & Clench, 1934c. 

Soft anatomy. Alcohol preserved specimens available 
29/dissected 17. Tanzania: BMNH 1/1, NHMW (no.47996) 
1/1. Malawi: CMNH 13/2, HM 5/5 (all now at RMNH), NG 
4/3 (2 now at BMNH), RMNH 3/3. Zimbabwe: NM 2/2. 
Additional Malawi specimens are in collections HM and NG. 
No others are known. Most specimens examined were well 
extended from their shells. 

In the individual specimen, the body colour varies from 
unicolorous pale buff to dark grey. A black thin-lined, 
coarsely reticulate pattern is characteristically present later- 
ally on the foot. Sole of foot is uniformly pale dusky, without 
variation in texture. Usually a delineated or diffuse narrow 
dark grey or black stripe, originating behind each ommato- 
phore, passes posteriorly toward the mantle on either side of 
the paler neck region. The mantle varies from unicolorous 
dark or light grey to a grossly maculate pattern. The Chirinda 



Forest, Mount Selinda, Zimbabwe field notes of A.C. van 
Bruggen in lift. ) record the body colour varying from black to 
pale grey marbled with black, and the longitudinal stripes 
varying from white to greyish white. 

The short, slender penis (P) and the slightly shorter penis 
sheath (PS) seem diminutive compared to the relatively gross 
structures of the basal female conduit. Of the 17 specimens 
dissected, 11 had the apical-most P and the basal-most basal 
vas deferens (BVD) projecting slightly above the rim of the 
PS. In the other 6 specimens, the P was completely covered 
by the PS, 2 of which were gravid, 2 were over-drowned with 
consequent eversion of the genital atrium (GA), one was 
immature and one was severely distorted because of 
improper fixation. There was no positive correlation in this 
latter group of 6 with such other possible influencing factors 
as latitude, shell size, length of penial retractor, month of 
collection or size of albumen gland. The specimen (NM) 
depicted in Fig. 17 collected by A.C. & W.H. van Bruggen in 
Zimbabwe (Vumba Circular Drive, Zonwi River Bridge) has 
thus been selected as representative of the first group and 
typical of this species. 

In all specimens, the penial retractor (PR) inserts either on 
the anterior diaphragm, on the body wall of the neck, or at 
the forward junction of the diaphragm, mantle and body wall. 
In well extended specimens, it can appear inordinately long. 
Cutting and spreading the thin-walled PS reveals the slender 
P and BVD within (Fig. 18). Fine muscle strands pass basally 
from the PR to cover thinly the apical P. About half way 
down the P, these strands attach to the smooth, shiny inner 
surface of the PS and then proliferate into voluminous strands 
that completely cover the BVD and infuse with the tissues of 
the basal P and PS. At this level the tissue layers are not 
distinct, but below the PS the male conduit continues as a 
short penial atrium (PA) that joins the vaginal atrium (VA). 
These fuse to form the genital atrium (GA). Strap-like 
overlapping, glistening eversion muscle bands (EM) connect 
the basal male and female conduits to the inner right body 
wall. During precopulatory behaviour, contraction of these 
muscles cause the GA, then the PA and VA to evert, to 
protrude as a stimulatory organ, and subsequently to initiate 
the extroversion of the intromittent organ. In some older 
specimens, these bands can be so voluminous and so high on 
the PS that they seriously obscure relationships. The thin- 
walled BVD passes through the PS to emerge as a much 
larger, thick-walled apical vas deferens (AVD), which basally 
functions as an ejaculatory duct and as a support for the 
thin-walled everted P. About midway apically, the AVD 
gradually becomes thin-walled until at the AVD/FO junction, 
it forms a glandular funnel-shaped chamber that internally is 
crowded with extremely thin epithelial partitions. This may 
function as a secondary seminal vesicle (SSV). There is no 
pilaster or verge in this species. 

In the basal female conduit, the vagina (V), free oviduct 
(FO) and the spermathecal duct (SD) form, without distinct 
delineations, an impressive muscular Y-shaped structure 
(Fig. 18). Upon dissection, the broad lumen and the promi- 
nent longitudinal plicae of the V are seen to be uninterrupt- 
edly confluent with those of the SD. In contrast, the lumen of 
the basal, muscular thick-walled FO is slender and it is 
confluent with that of the V through a small recessed pore. 
This arrangement ensures the passage of the intromittent 
organ into the SD during copulation. The thick-walled FO 
functions basally as an ovijector; apically its thin walls 
accommodate the descending eggs. The clavate-elongate 



28 

spermatheca (S) (Fig. 17) frequently is apically attenuated 
because thin fibers firmly attach it 4-5 mm below the 
AVD/FO junction. That junction, in turn, is held tightly 
together on the right internal body wall by fibres from the 
transverse myoseptum. Any contraction or extension thus 
provides maximum pull in this part of the body. As a result, 
under certain conditions, the S apex and SD may become 
inordinately elongate or nearly disappear into a huge saccu- 
late S. Severe contraction of the viscera may even produce 
the artifact of a tandem bilobed S. The only important 
character of the S is its consistent position below the 
AVD/FO junction. The FO, though shorter than the AVD, is 
conspicuously long, nearly uniformly wide, and approxi- 
mately twice as wide as the AVD. Although there are thin 
facia binding together V, SD, S, FO and AVD, there is no 
formation of a vaginal rententor per se. 

Two specimens were gravid. One from Chinyama, Malawi 
(HM) was collected in February 1983 (mid-rainy season) and 
had 56 light yellow, moderately large eggs averaging 
6.3 x 5.2 mm. The other, from Lake Rukwa, Tanzania 
(BMNH), was collected in 1938 (month not indicated) and 
had 51 similar eggs measuring 7 x 5.5-6.5 mm. The eggs, 
without discernible embryos, were closely embraced by uter- 
ine tissue folds as depicted by Mead (1950, fig. 48) for 
Achatina fulica Bowdich, 1822. 

All specimens dissected showed a remarkably gross devel- 
opment of a trimerous hermaphroditic duct system (Fig. 19). 
Basally, there is a 10 mm characteristically deeply convoluted 
portion just distal to the talon. Next, there is a 3 x 2 mm 
discreet, compacted saccular portion that appears to be 
glandular. And finally, there is a 6 mm slender, weakly 
convoluted portion that quickly forms a series of five larger, 
tightly convoluted ductules leading to the five gonadal acini 
buried in the right lobe of the digestive gland. The talon is 
elongate, capitate and diminutive (1.5 x 0.5 mm). The sper- 
moviduct is characteristic of the family. 

The following anatomical characters distinguish this species 
from B. graueri: basal genital fascia diaphanous; AVD about 
twice the width of BVD and half the width of FO; P slender, 
much longer than wide, normally projecting slightly above 
PS; BVD slender, much longer than wide. 

Type material. For over a century, Bourguignat's (1889) 
Serpaea pintoi has been an engima to conchologists. The 
principal contributing factor has been the artist's rendering an 
excessively bold, wide, broadly truncated columella. A sec- 
ond factor is that the exceedingly thin, fragile shell of the only 
know specimen unfortunately had become broken sometime 
since it was drawn. The several pieces, including a figure '4' 
label, had been placed in a separate vial. This specimen (Figs. 
41, 42; Table 8) is in Paris (MNHN) and is here considered to 
be the holotype by monotypy (Code Art. 73(a)(ii); 74(b). Its 
conchological features and the type locality in eastern Tanza- 
nia support the conviction that this is conspecific with Smith's 
junior subjective synonym Achatina fragilis (1899). Pilsbry 
(1904:1, 21, 63) placed Serpaea in the synonymy of Achatina, 
indicated (1909:113) that Smith's name was a primary hom- 
onym of Deshayes' fossil species (1864), proposed the 
replacement name Achatina nyikaensis, reproduced Smith's 
figures [fig. 26 is way too intensely coloured], and perspica- 
ciously placed Bourguignat's species and Smith's species 
seriatim under Achatina in his Manual. 

Smith's syntype lot of seven specimens from Nyika Plateau, 
Malawi, unfortunately contains a single specimen of Achatina 



A.R. MEAD 
Table 8 B. pintoi - Representative shells measurements. 







Greates 


t Apert 


are 


Last 


/o 






Whorls 


Length 


Width 


Length Width whorl LW/L 


. % 


W/L 


6V2 


74.6 


42.8 


47.6 


26.1 


62.5 


84 


57 


Nyika 
Plateau 
(BMNH) 
Lect A. 
fragilis* 


6>A 


74.4 


42.0 


46.3 


24.6 


61.4 


82 


56 


Macequese 
(SAM) 


6 


66.8 


40.6 


46.9 


24.0 


58.0 


87 


61 


Nyam- 
badwe 

(Nor 


6V4 


62.2 


32.7 


38.3 


19.5 


50.6 


81 


52 


Utengule 
(ZMB) 
Holo C. 
thielei * 


6 


56.2 


37.8 


38.0 


21.3 


47.9 


85 


67 


Bulawayo 
(RMS) 


6 


56.0 


33.0 


35.5 


19.6 


46.5 


83 


59 


Utengule 
(BMNH)* 
Mac 
Andrew 

(1563) 


6 


53.3 


27.2 


33.0 


16.7 


43.5 


82 


51 


Chirinda 

(BMNH)* 

1907.7. 

25.3 


6 


50.1 


32.6 


33.0 


19.8 


42.4 


85 


65 


Mamboya 
(BMNH)* 
1885.5. 

25.47 


6 


49.7 


33.6 


31.8 


19.5 


43.2 


87 


68 


Ngrengre 
(MNHN) 
Holo S. 
pintoi* 


6'/4 


49.3 


27.2 


28.3 


15.0 


38.5 


78 


55 


Tanzania 

(NHMW)' 


5% 


49.3 


33.7 


34.9 


19.3 


42.7 


87 


68 


Pemba 
(SAM) 


5% 


43.7 


29.0 


29.0 


17.8 


37.2 


85 


66 


Usagara 
(BMNH) 



Total specimens examined: 92. Sources: BMNH. FMNH, HM. 1RSN. LNK. 
MCZ, MNHN. MRAC, NG. NHMW. NM, NMW. RMNH. SAM. SMF. 
UMMZ, ZMB. 

craveni Smith, 1881. Since Smith, as its author, was thor- 
oughly familiar with this latter species, since he did not 
designate the number of syntypes, and since he separately 
discussed (p. 35) and figured (figs. 1-4) specimens of both 
this species (BMNH no.97. 12.31. 1-7) and A fragilis (BMNH 
no. 97. 12.31.8-14) in his 1899 paper, it is here safely assumed 
that the division between the two adjacent accessioned speci- 
men lots was an inadvertent curatorial error, with no nomen- 
clatural implications for the misplaced specimen. The 
unicolorous, largest and finest specimen (BMNH 
no. 97. 12. 31. 9) of the six syntypes (Smith's fig. 3, Pilsbry's fig. 
25), here shown in Figs. 43, 44 bears a handwritten note, 
'Lectotype "3" A.C. van Bruggen, May 1974'. This selection 
is here endorsed. Other known and examined paralectotypes 
are single specimens in Berlin (ZMB no. 101934) and Vienna 
(NHMW/R). 

Ancey (1902) described and illustrated a specimen which 
he labelled "Achatina . . . sp. nov. ' from Ugogo (5° 4' S, 34° 
4' E). Ancey's collection was distributed widely by Geret and 
this specimen so far has not been located. However, the very 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



29 



large aperture, the slender arcuate columella, the transverse 
stripes, the translucence of the shell showing the pattern 
through the aperture, and the locality indicate that his 
specimen is Bourguignat's species. 

Bequaert & Clench (1934c) found nine purchased (Her- 
mann Rolle) specimens from Utengule, Tanzania in the 
Berlin Museum (ZMB) that were marked as new and given a 
preempted manuscript name, but were undescribed. 
Although they felt the new taxon 'might perhaps prove to be 
a local race' of Callistoplepa nyikaenisis , its more slender, 
more tapered shape convinced them it should be established 
as the new species Callistoplepa thielei. The largest specimen 
previously had been broken, so they selected the second 
largest specimen as the holotype (Figs. 45, 46). In addition to 
the holotype, they figured two paratypes. This series of four 
types (ZMB no. 53177) are in Berlin (Kilias, 1992). Examina- 
tion of these types in Berlin in 1989, showed evidence that the 
paratypes had been broken apparently on their return to 
Berlin because their fragile shells had been stuffed exces- 
sively with cotton, leaving only the holotype undamaged. The 
five other original specimens, 3 full grown and 2 juveniles, 
are labeled paratypes (MCZ no. 98686) in the Harvard collec- 
tion. Three other broken specimens with the same data are in 
Frankfurt (SMF); there is no evidence that these were seen 
by Bequaert & Clench. 

Type locality. Bourguignat (1889) states,' . . . provient 
des environs de l'Ougerengere, vallee du Kyngani, dans 
l'Oukani'. Inside the lip of the holotype, in Bourguignat's 
handwriting, is 'Serpaea pintoi Ougerengere (Oukami). A 
small label bears the inscription 'M. Requin 1846 30'. 
According to B. Verdcourt (in litt.), Ukami is a large 
geographic district that surrounds and includes the Uluguru 
Mountains in Tanzania. The Kingoni river and the Ngeren- 
gere settlement and stream, however, place the type locality 
close to 7° 03' S, 38° 31' E, 125 km west of Dar-es-Salaam (cf 
Verdcourt, 1966:111). 

Distribution. This is by far the most widespread species in 
the genus. The 32 recorded localities define a 1800 x 1200 
km territory 4-20° S, 27-39° E that includes eastern Zaire, 
eastern and western central Tanzania, nearly all of Malawi, 
south central Zambia, southern and eastern Zimbabwe, and 
far west central Mozambique (Fig. 16). The northern outpost 
of this species was established in Zaire by R. Grauer in 1910, 
when he found and preserved in alcohol a specimen in 'the 
primary forest behind bordering hills of the northwest shores 
of Lake Tanganyika, 1800-2200 m' (trans. ) 4° 30' S, 29° 00' E. 
It is noteworthy that along the west, there is an almost 
straight N-S line of demarkation 27-28° E from eastern 
central Zaire to south central Zambia and southwestern 
Zimbabwe. Connolly, in his writings (1925, 1939) and on 
some of his specimen labels, juxtaposes the geographic names 
Macequece (a district in eastern central Mozambique) and 
Lourengo Marques (the major port now known as Maputo in 
southern Mozambique). Specimens may have been shipped 
from this port, but there is no convincing evidence that this 
species has ever been collected in Mozambique south of the 
Macequece district. The remarks of Germain (1935:4) seem 
to clarify when he explains that Portuguese East Africa is 
generally divided into two large regions separated by the 
Zambeze River, 'le Mozambique au Nord, le Lourenzo 
Marques au Sud'. When adequate population studies can be 



made in this widespread species, valid subspecies may 
emerge. 

Remarks. This apomorphic species is most closely related to 
B. marteli. Both Achatina craveni E.A. Smith, 1881 and B. 
pintoi are highly variable and are often confused in collec- 
tions, particularly where the field data are the same or the 
individuals are small. Despite rather considerable overlap in 
the extremes of the conchological characters of these two 
species, the shell of A. craveni can be differentiated on the 
basis of the following: nepionic whorls smooth; last whorl 
equals only 70-75% of shell length (vs 80-90%); one to two 
more whorls for the same length; apex more acute; columella 
much shorter, broader, straighter and more squarely trun- 
cated; finer and deeper granulate-cancellate sculpture; shell 
usually much less fragile. 

The wide distribution and the independently great variabil- 
ity in shell characters, even within a single population, are 
responsible for the long and confused synonymy of this 
species. The soft anatomies of antipodal specimens, and 
many between, support the conclusion that only one species 
is involved. 

Ecological notations with specimen data (HM, NG) indi- 
cate that active specimens were found in Malawi lowland 
evergreen forests, along the banks of earth roads, crawling on 
leaf litter, and on the underside of banana leaves in the rain. 
Specimens from Mbeya, Tanzania were found in luxuriant 
herbaceous vegetation (LNK). The director of the Imperial 
Institute of Entomology earlier reported that in the Nkota- 
Kota district of Malawi there was a 'very serious outbreak in 
November 1937', implying that this species has the potential 
under certain conditions of becoming an agricultural pest. In 
Zambia this snail is know as 'chuzuya'. 

The fine series of specimens in BMNH convincingly dem- 
onstrates the wide range of variability in the shells of this 
species (Figs. 41-51). 

Bequaertina fraterculus (Dupuis & Putzeys, 1900) 
Figs. 52, 53 

Ganomidos fraterculus 

Dupuis & Putzeys, 1900:xiii, text fig. 18. 
Callistoplepa fraterculus 

Pilsbry, 1905:129, pi. 47, fig. 23 (ex Dupuis & Putzeys); 

Germain, 1909:90; Pilsbry, 1919:80; Bequaert & Clench, 

1934c: 114. 

Shell. Shell ovate-turrite, extremely thin, translucent. 
Whorls 6-6'/4. The second and third whorls are comparatively 
large, long and nearly straight-sided, producing characteristi- 
cally a collared blunt mammillate apex. The fourth and 
subsequent whorls are convex and expanded proportionately. 
Sutures moderately deep. Last whorl large, but not inordi- 
nately so, 78% of shell length; range for 6-6'A whorls, 
77-80% (n = 10). Aperture oval, pale milky within. Col- 
umella brown, long, slender, nearly straight, obliquely trun- 
cated. Outer lip thin, evenly arcuate; receding at base in 
profile. Parietal callus diaphanous. 

Highly irregular castaneous streak and spot brush marks, 
some of which are closely highlighted with buff adaperturally, 
are characteristically found on the last whorl; however, these 
may be reduced to a few obscure dull buff spots more or less 
limited to the peripheral carina. The earliest signs of this 
diagnostic colour pattern are seen in the third whorl. The 
ground colour intergrades from pale horn colour of the 



30 

unicolorous nepionic whorls to an obscuring dark olive brown 
of the last whorl. 

Faint, closely oppressed minute crescentic granulations 
may appear in the second half of the otherwise smooth first 
whorl. These granulations dominate the second whorl, giving 
it an evenly, delicately engraved appearance. In contrast, 
rather bold, diagonally oriented arcuate plicae, arising from 
the suture below, disrupt this sculpture in a narrow basal 
zone. This plication, which may start even in the last part of 
the first whorl, rather abruptly disappears in the third whorl, 
leaving a delicate, closely aligned series of nearly transverse 
lirae, uninterrupted except for small, sparce, ghost-like 
patches of granulations in some specimens. This marks the 
end of the nepionic whorls. In the fourth and fifth whorls, the 
lirae become increasingly strongly prosocline, more regular, 
apically arcuate immediately below the suture, and eventu- 
ally so prominent that they obscure the growth lines. In the 
sixth whorl, the lirae become finer, less regular, more nearly 
orthocline, and often interspersed with short parallel lirellae. 
In the fifth whorl, spiral lirae, which seem to join rather than 
interrupt the prosocline lirae, appear somewhat irregularly, 
producing a slight checkerboard effect, reminiscent of the 
sculpture of Achatina tracheia Connolly, 1929. These spiral 
lirae, which are quite close together near the suture, irregu- 
larly diminish in number and intensity toward the columella. 
Very shallow malleations, starting subtly in the fourth whorl, 
intensify the checkerboard effect. A subdued but apparent 
carina appears at the periphery, where the lirae are seen to 
bend slightly. The carina is more conspicuous in the younger 
specimens. Abrasion of the shell along the periphery and 
along some of the more prominent lirae visually intensifies 
the carina and the sculpture by exposing the brilliantly shiny 
inner periostracal layer. The outer layer of the periostracum 
imparts a characteristically dull, corneous luster to the shell. 

Soft anatomy. No known alcohol preserved specimens. 

Type material. Nine of the ten known specimens of this 
rare species were collected by Dupuis and are considered 
syntypes. Six of these were identified and labelled as 
'cotypes': three in Tervuren (MRAC no. 5140-5142) and 
three unnumbered specimens in Bruxelles (IRSN, General 
Collection). The seventh specimen (MRAC no. 5139) was 
labelled as the one figured by Dupuis & Putzeys (1900); their 
figure 18 (reproduced by Pilsbry, 1905) is so generalized that 
it cannot specifically identify with any of the syntypes. 
Unfortunately, the apex of this specimen had been broken 
and cemented together, with a resultant alteration of the shell 
configuration and length. The damage possibly occurred 
during the precarious period when the artist had the speci- 
men. Two additional syntype specimens (IRSN) were given 
to Dauzenberg by Dupuis, one of which was collected in 
Nsendwe. Under these circumstances, and since the authors 
did not designate a holotype, the largest and finest of the four 
MRAC syntypes (no. 5140) is here selected as the lectotype of 
Dupuis & Putzeys' Ganomidos fraterculus (Figs. 52, 53; 
Table 9). The tenth known specimen, acquired by Preston, 
was passed on to V.W. MacAndrew, and is now in the 
BMNH. Its apex was damaged and repaired naturally. This, 
too, most probably was collected by Dupuis and should be 
considered a paralectotype. 

Type locality. The Island of Mvula on the Lualaba 
River, Zaire. J.C. Bequaert was unable to find this locality 
on any map (Pilsbry, 1919:11, 19), nor is it listed in the 



A.R. MEAD 
Table 9 B. fraterculus - Representative shells measurements. 



Whorls 


Length 


Greates 
Width 


t Aperture Last % 
Length Width whorl LW/L 


. % 


W/L 


6 


51.0 


28.7 


30.0 


16.4 


40.7 80 


56 


Mvula 

(BMNH) 

PLec 


6 


50.5 


29.0 


30.0 


16.9 


39.3 78 


57 


Mvula 

(IRSN) 

PLec 


6V4 


50.4 


28.5 


30.5 


17.0 


38.9 77 


56 


Mvula 
(MRAC) 
5140 Lect* 


6 


47.5 


27.4 


27.8 


16.0 


37.5 79 


58 


Mvula 

(MRAC) 

5139 


6 


46.3 


26.8 


28.7 


15.5 


36.7 79 


58 


Mvula 
(MRAC) 
5141 PLec 


6 


44.4 


26.6 


27.2 


14.8 


35.0 79 


60 


Nsendwe 

(IRSN) 

PLec 


6 


44.3 


26.4 


26.7 


14.5 


34.0 77 


59 


Mvula 

(IRSN) 

PLec 


5% 


40.2 


26.4 


26.2 


14.2 


32.6 81 


66 


Mvula 
(MRAC) 
5142 PLec 


5% 


37.5 


23.0 


22.6 


13.2 


29.7 79 


61 


Mvula 

(IRSN) 

PLec 


5% 


36.5 


21.7 


22.0 


12.2 


28.2 77 


59 


Mvula 

(IRSN) 

PLec 



Total specimens examined: 10. Sources: BMNH. IRSN. MRAC. 

current USBGN series. The only lead is the fact that a 
single specimen of this species in the Dautzenberg collec- 
tion (IRSN) bears the data: 'Nsendwe, Congo. P. Dupuis 
Coll., leg & ded\ At the same time that Dupuis & Putzeys 
described Ganomidos fraterculus (1900), they described 
the new species Perideriopsis mvulaensis, giving its locality 
as 'ile de Mvula (en face de Nsendwe)'. Bequaert in Pilsbry 
lists Nsendwe as 3° 05' S, 26° E. This location nearly 
coincides with the important crossroad Kindu-Port-Empain 
2° 57' S, 25° 56' E, hence Mvula must be very close to 3° S, 
26° E. 

Distribution. The two known localities for this species are 
the Island of Mvula and the nearby onshore village of 
Nsendwe in Zaire. Given the nature of the riverine environ- 
ment, this species may well be found on other of the many 
small islands and possibly in shore sites along this northward 
flowing remote section of the Lualaba. 

Remarks. Phylogenetically, this species appears to stand 
between B. graueri and the plesiomorphic B. pellucida. 
Based on a single specimen from Nsendwe (BMNH, Preston 
'L/K 13/11/01'), B. pellucida may be sympatric with B. 
fraterculus. 

Bequaertina graueri (Thiele, 1911) 
Figs. 54-57 

Achatina graueri 

Thiele, 1911:205, pi. 5, fig. 43; Pilsbry, 1919:78. 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



31 



Achatina (Cochlitoma) graueri 

Pilsbry & Cockerell, 1933:366, pi. 1, fig., 1, la. 
Callistoplepa graueri 

Bequaert & Clench, 1934c:115. 
Callistoplepa graueri 

Schouteden, 1935a:110; 1935b:287; Oliver, 1983:9. 
Callistoplepa babaulti 

Germain, 1936:151, text fig. 46. 

Shell. Shell lacrimoid-subsuccineiform; the thick, durable 
periostracum appears to provide more support than the thin 
calcareous shell. Whorls 6-6V2, rarely 7. Spire tapered, 
mammillate, elevated-conic, clearly shorter than aperture 
length. The first IV2 whorls form a bluntly obtuse dome. The 
second and third whorls expand only slightly, but descend 
rapidly; this produces the strongly mammillate apex. The 
fourth and following whorls descend proportionately, are 
increasingly more convex, but expand rapidly to produce an 
inordinately large last whorl. Sutures moderately deep, 
increasingly so between fifth and sixth whorls. Last whorl 
80% of shell length; range for 5-7 whorls, 77-83% (n = 50). 
Aperture oval to elongate; very thin blue-white shelly layer 
within. Columella markedly slender, almost entirely concol- 
orous, slightly arcuate or nearly straight; typically very nar- 
rowly and obliquely truncate. Outer lip thin, usually 
somewhat obscured by the more rapidly advancing thick 
periostracum, which tends to curl into the aperture in dried 
specimens. The arc of the outer lip is often greatest below 
midway and basally extended well below the columellar 
truncation in the more mature specimens. Parietal callus 
appears to be virtually absent in the smaller specimens and 
barely visible in the larger and older specimens. 

The first three whorls are unicolorous beige-buff to buff- 
horn colour. Faint, diffuse wide transverse castaneous bands 
may emerge in the second whorl and become darker, more 
conspicuous and fragmented in the third and fourth whorls. 
These gradually give way to narrow transverse bands of 
various shades of brown, irregularly appearing coincident or 
alternating with growth bands. The darkest, broadest bands 
usually indicate interrupted growth. In some specimens the 
transverse bands may be essentially absent. Ground colour 
varies within and between specimens in a spectrum of olive- 
buff, light yellow-brown, olivaceous brown and medium dark 
brown. The fifth whorl often has a contrastingly paler ground 
colour; hence, a 5-whorl specimen, with its short last whorl, 
may appear to be a different species. 

Pronounced spiral striae, usually 5-7, starting in mid-first 
whorl are offset by beaded or semilunar granules that are 
irregular in size and not aligned in vertical rows. The spiral 
rows of granules increase to 12-17 or more and the now very 
small granules become more uniform, more prominent, and 
gradually more transversely aligned in the second and third 
whorls. At the end of the third whorl, there is a prominent 
delineation that marks the end of the nepionic whorls. At this 
point, the granulate sculpture is rather abruptly taken over by 
thin, slightly prosocline growth wrinkles that increase in size 
and number until they dominate in the fourth whorl. In the 
fifth whorl, there is a reemergence of the granulate sculpture 
in the form of notched, subquadrate, tile-like plates that 
often resemble the block letters K,H,W,V,Y & M, as is 
strikingly seen in Achatina reticulata Pfeiffer, 1845. This 
reappearance of the granulate sculpture, accentuated by the 
deeper spiral striae, dominates the fifth whorl, although the 
growth wrinkles continue to increase in calibre and become 



subcrenulate apically. In the usually somewhat darker 
coloured last whorl, the granulate sculpture is once again 
greatly reduced, but may be highlighted here and there by an 
isolated sporadic deep section of a spiral stria. The earliest 
malleations appear subtly in the third or fourth whorl and 
intensify in the following whorls. They consist of usually short 
spiral or diagonal ridges that join or distort the growth 
wrinkles to form a coarse, irregular raised network of welts. 
Often entering into this is a very faint elevation at the 
periphery, below which the otherwise nearly uniform sculp- 
ture is reduced. An occasional specimen is entirely without 
malleations. Their irregular appearance is probably explained 
by thin shell, tough perostracum, and environmental impacts. 
Continued shell deposition from within 'fixes' the dents in 
place. Rarely is the outer layer of the thick periostracum 
broken enough to reveal the shiny inner layer. 

Soft anatomy. Alcohol preserved specimens available 
82/dissected 9. Zaire: MRAC (Mulungo, no. 204. 632-633) 
2/2; (Kahusi-Tshibati no. 610.302-305, 342-343) 79/6; ZMB 
(lectotype) 1/1. Unfortunately, all dissected specimens except 
the lectotype, were exposed excessively to formalin during 
their preservation; thus, even with prolonged special treat- 
ment, their tissues remained hard and the specimens were 
exceedingly difficult to dissect. Pilsbry and Cockerell (1933) 
described the living animal as very pale ochreous with head 
and broad based tentacles faintly bluish. 

Clearly, the most conspicuous and characteristic aspect of 
the basal genital conduits is the vaginal retentor (VR) muscle 
system, grossly dominating in ventral view (Fig. 20). Slender, 
parallel, glistening, partly fused muscle bands pass ventrally 
from the vagina (V) to the right body wall along a dorsolat- 
eral line from immediately posterior to the genital aperture to 
the junction of the mantle and the right body wall. From the 
left side of the vagina, this system gives rise to a series of 
muscle bands that starts at the peniovaginal angle and binds 
tightly together the equally prominent apical vas deferens 
(AVD) and the free oviduct (FO). A more bold series of 
distinct, but laterally fused muscle bands pass from the left 
lateral aspect of the AVD to the same right dorsolateral line 
of attachment, further obscuring the basal genital structures. 
(These bands are shown cut short in this figure.) Apically this 
whole muscle system is reduced largely to a thin, transparent 
membrane attached to the junction of the AVD and the FO. 
At this same junction, a similar membrane passes to the 
spermatheca (S) and attaches it to the FO. The S is propor- 
tionately extremely large in this species. In four of the nine 
specimens examined, the slender apex of the S is folded back 
on itself, as shown here; this is apparently a common artifact 
of preservation. In the rest, the apex is extended, and 
although it may appear to go apically beyond the junction of 
the AVD and FO, it is not attached to the spermoviduct (SO) 
as it is in many achatinids. The spermathecal duct (SD) is so 
short and broad that when the S is gorged, it seems to be 
sessile on the V. 

Short, often diagonal or anastomosing muscle bands, sepa- 
rate or combined, are found at the base of the penis sheath 
(PS) and the V. These eversion muscle bands (EM) initiate 
the precopulatory extroversion of the genital atrium (GA). 
The PS normally completely envelops the short, diminutive 
penis (P); in only one of nine specimens, the apical P 
projected slightly. The penial retractor (PR) has its origin on 
the apical P and inserts on the mid-forward diaphragm at or 
near the junction with the body wall. A single specimen, 



32 



A.R. MEAD 



shown here, had a bifurcate insertion; a multiple insertion is 
not rare in the achatinids. At the origin of the PR, the muscle 
fibrils enshroud smoothly and completely the apical P and the 
robust basal vas deferens (BVD), greatly obscuring the 
relationships in the basal male conduit (Fig. 21). Contributing 
to this, the PS is free from the P only in the approximate 
upper half of the left side. On the right side, fibrils from the 
PR extend basally to form a dense webbing that seems to 
invade the substance of the outer wall of the P and the inner 
wall of the PS. Below the PS, this infusion of tissues, along 
with the EM, obscurely defines the wall of the penial atrium. 
This atrium connects the lumina of the P and GA (Fig. 22). 
The lumen of the P is thickly carpeted with vermiculate 
rugae, which become slender and elongate near the GA, 
resembling the plicae of the basal V. No verge or pilaster is 
present. Eccentrically in the apex of the P, a small aperture 
leads to the narrow lumen of the extremely thick-walled BVD 
and AVD. These two structures provide the physical support 
for the extroverted, highly expansile P. Since they have 
supportive and ejaculatory functions, they may explain the 
extreme development of the VR. Approximately 8-10 mm 
basal to the junction of the AVD and FO, the lumen of the 
AVD enlarges considerably and forms an elongate, thin- 
walled chamber, which conceivably functions as a secondary 
seminal vesicle (SSV). Apically this chamber becomes saccu- 
lar with thin elongate rugae. 

A single specimen (610.343) was gravid. Six large, fully 
formed eggs, 9.5 x 6.5 mm, were in the apical (oviductal), 
cream coloured portion of the spermoviduct; no eggs were in 
the contrastingly light brown uterine basal portion. This 
specimen and the five others in the same lot had robust, 
mature coloured, fully formed reproductive tracts. The field 
data thus indicate that in the Kivu, breeding takes place in 
October. 

Kidney is large, typical of the subfamily, broad anteriorly 
and truncated posteriorly. Five ovotestis acini are embedded 
in the columellar surface of the right (apical) lobe of the 
digestive gland. The inconspicuous anterior aorta is on the 
left posterior surface of the lung, where it penetrates the 
diaphragm. The hermaphroditic duct, similar to that of B. 
pintoi (Fig. 19), is trimerous with an abruptly enlarged 
saccular central portion, 5.3 x 2.4 mm. 

The following anatomical characters distinguish this species 
from B. pintoi: basal genital fascia gross, forming a VR with a 
massive system of muscle bands; AVD, BVD and FO are all 
about the same width; P is strikingly short and stubby, 
normally retained entirely or nearly entirely within the PS; 
BVD wide, about as long as wide. 

Type material. Thiele (1911) described this species from a 
single mature specimen (ZMB no. 101937) in the Schubotz 
collection and two small juvenile specimens from the Grauer 
collection. The mature specimen is nearly full grown but only 
moderately large (Figs. 54, 55; Table 10). Thiele's fine line 
illustration shows it slightly larger than natural size in aper- 
tural view only. This specimen, whose soft parts are in 
alcohol, I labelled as the lectotype of Thiele's Achatina 
graueri when I examined it in East Berlin (ZMB) in August 
1989. At that time, the single available very dark coloured 
juvenile specimen (Table 10) therefore was labelled paralec- 
totype. Since then a second small paralectotype has been 
found there and so labelled by Kilias (1992). 

Germain (1936), without any apparent knowledge of 
Thiele's species, described and figured the junior subjective 



synonym Callistoplepa babaulti from two specimens collected 
by Babault in Kitembo, Kivu, comparing it only with C. 
marteli. The two Paris syntypes (MNHN) are large, typical 
specimens of Bequaertina graueri (Table 10). The larger, 
finer specimen is here selected as the lectotype of Germain's 
C. babaulti (Figs. 56, 57). 

Type locality. Idjwi Island (= Kwidschwi, Kwidjwi, 
Idjewi), Lake Kivu, Zaire 2° 09' S, 29° 04' E. 

Distribution. This species occupies a 450 km long, narrow, 
north-south corridor in the upper Rift Valley of Zaire 
between Beni and Uvira (Fig. 16). So far, it has been found 
only as far west as Kitembo and projects slightly east of Zaire 
into Lobengera Mission, Rwanda and into Ibanda, Uganda. 
Eventually, it will also be found in Burundi. 

Remarks. This is the largest and most distinctive species in 
the genus. It is most closely related to Bequaertina fratercu- 
lus. Because of its size and colour, and because in some 
localities it is sympatric with Achatina stuhlmanni von Mar- 
tens, 1892, it has been confused with that species. However, 
since A. stuhlmanni has a shorter, broader spire, a more 
obtuse apex, and a distinctive spirally fine-combed wavy 
sculpture (Bequaert & Clench, 1934a:3), it can readily be 

Table 10 B. graurei - Representative shells measurements. 



Greatest Aperture Last % 
Whorls Length Width Length Width whorl LW/L % W/L 



7 93.2 


45.7 


52.3 


26.6 


71.8 


77 


49 


Katana 

(MRAC) 
610.304 


6'/4 89.7 


47.0 


55.5 


28.4 


72.6 


81 


52 Tshibinda 
















(MRAC) 

5115 


7 89.3 


43.6 


50.7 


25.5 


70.0 


78 


49 


Katana 

(MRAC) 

610.302 


6'/4 78.4 


42.5 


46.8 


24.3 


62.5 


80 


54 


Kitembo 
(MNHN) 
Lect C. 
babaulti* 


6 77.0 


38.6 


45.9 


23.5 


63.0 


82 


50 


Uvira 

(MRAC) 

607.170 


6>/ 4 71.8 


41.4 


41.9 


23.4 


56.1 


78 


58 


Kitembo 
(MNHN) 
PLec C. 
babaulti 


6 62.0 


36.5 


37.8 


20.6 


49.1 


79 


59 


Idjwi 
(ZMB) 
101935 
Lect A. 


5% 49.8 


29.1 


31.8 


16.4 


40.0 


80 


58 


grauer?* 
Idjwi 


6 49.2 


22.8 


31.0 


14.5 


39.5 


80 


46 


(MCZ) 
Beni 
(MRAC) 
5119 


5V4 42.0 


25.7 


27.0 


14.0 


33.0 


78 


61 


Idjwi 
(ZMB) 
101936 
PLec A 

graueri 


Total specimens 


examined: 


54. Sources: IRSN 


I,MCZ 


, MNHN, MRAC, 


NMW, RMNH, 


SMNH, UMMZ, ZMB. 











NEW SUBFAMILY AND GENUS ACHATINIDAE 



33 



distinguished. Pilbry & Cockerel (1933) reported seeing two 
living specimens 'crawling about 5 feet up on the trunk of a 
tree in the forest above Tshibinda at about 2100 m' in Zaire. 
The largest collection of this species, including many alcohol 
specimens, is to be found in Tervuren (MRAC). B. graueri is 
the type species of the genus. 

Radulae and jaws 

D'Ailly (1896:69) was the first to describe and illustrate the 
radulae of Callistoplepa shuttleworthi and C. barriana. Pilsbry 
(1904:ix, xv) referred to d'Ailly's work but reproduced only 
the illustration of the latter species. He also reported (p. 72) 
that G. Schacko (1881) {nee 'Schako 1 ) found 'A pulchella has 
... a very small central tooth'. The present work shows 
Schacko's specimen was therefore misidentified. Thiele 
(1929:560) examined and illustrated in part the radula of C. 
shuttleworthi. He also examined but did not illustrate the 
radula of Leptocala mollicella and pointed out that the middle 
tooth was a little smaller than the neighbouring teeth. Possi- 
bly on the basis of this observation, he prophetically juxta- 
posed 'Callistoplepa and Leptocala. More recently, Ortiz & 
Ortiz (1959:47) also illustrate the radula of C. shuttleworthi, 
but the focal plane of the microscope was apparently too low 
and the configurations of the teeth are misleading. The 
radulae in the present project were prepared according to the 
recommendations of Solem ( 1972) and the emphasis has been 
placed on the rachidian teeth and the adjacent laterals. 

Because the soft anatomies of the four callistoplepine 
species are so similar, it was not surprising to find the radulae 
of C. barriana, C. shuttleworthi and L. mollicella (Figs. 
58-63) to be remarkably similar. This fact supported the 
earlier decision to conserve intact the odontophores of the 
two extant soft anatomy specimens of L. petitia. It is assumed 
with confidence that the radula of this latter species is 
essentially like the others. In those examined, all have bold 
rachidian basal plates and a broad functional rachidian tooth 
that is one-half to two-thirds the size of the adjacent lateral 
teeth. A second type of tooth is found in the first series of 
laterals, which similarly consists of broad, solitary meso- 
cones, but with conspicuous laterally asymmetrical basal 
plates. A third type of tooth arises in the second series of 
laterals, wherein the mesocones angle increasingly more 
mesad and small ectocones gradually arise. These merge 
almost imperceptibly into a fourth type, the tricuspid margin- 
als with minute irregular endocones, broad shorter serrate 
mesocones, and increasingly reduced basal plates that no 
longer contact the teeth posterior to them. The greatest 
irregularity within and between specimens occurs in the 
gradient between the bicuspid laterals and the tricuspid 
marginals. Hence the following formulae (tooth numbers 
from centre to right) are only approximate: C. barriana 
C-3 1-55-84, C. shuttleworthi C- 17-28-49, and L. mollicella 
C-19-28-66. 

The available radula specimens in only two of the five 
Bequaertina species have produced an incomplete and some- 
what confusing picture in this genus. The basal genital 
systems of B. pintoi and B. graueri are fundamentally similar 
- both reflecting affinities with the Zaire Basin subgenus 
Achatina {sensu Bequaert, 1950). It thus was anticipated that 
the radulae also would be similar. The radula of B. pintoi 
(Figs. 64, 65), not surprisingly, was found to be of the same 
type as that of Achatina craveni E.A. Smith, 1881 (Figs. 68, 
69). Both have greatly diminished, essentially nonfunctional 



rachidian teeth that are almost concealed by the adjacent 
laterals. And both have broad based, nearly tricuspid laterals 
with angular ectocones, broad mesocones and endoconal 
flanges. In addition, the mesocone column of each lateral 
tooth directly contacts and supports the broad basal plate of 
the tooth immediately posterior to it. It should be noted that 
A. craveni, on the basis of its soft anatomy, belongs in 
Bequaert's subgenus Achatina rather than where he has 
placed it in his subgenus Lissachatina. The surprise came in 
the radula of B. graueri (Figs. 66, 67), with its large functional 
rachidian tooth, attenuated massive basal plates, more 
restricted contact support between horizontal rows of teeth, 
and an imperceptible gradient into the marginal teeth. Within 
the genus, B. pintoi and B. graueri are at the conchological, 
geographic and ecological antipodes. The known plasticity in 
molluscan radulae suggests that undetermined different feed- 
ing demands in dissimilar habitats have produced the con- 
trasts in the radulae of these otherwise two closely related 
species. B. graueri and B. fraterculus appear to be very 
closely related conchologically. There is a question now 
whether the radulae will support this assumption. In reality, 
the relationships in Bequaertina will not be understood until 
both the soft anatomies and radulae of B. fraterculus, B. 
pellucida and B. marteli are known. Radula formulae: B. 
pintoi C-^2-25, B. graueri C-59, A. craveni C-34-24. 

The castaneous callistoplepine jaw forms an unusually 
broad middle section that quickly tapers on each side to about 
half its width and curves inward at the ends into a collariform 
structure. Its surface is featureless except for microscopic 
horizontal growth increments best seen under transmitted 
light. Measurements: C. barriana! x 1.3 mm, C. shuttlewor- 
thi 1.5 x 0.6 mm, L. mollicella 1.7 x 0.4 mm. The illustra- 
tion of Ortiz & Ortiz (1959:46) for C. shuttleworthi appears 
excessively broad. 

The jaw of B. pintoi forms a light castaneous nearly uniformly 
slender rainbow arc, 5.5 x 1.1 mm, with ca 36 irregularly placed 
vertical ridges. In B. graueri the jaw forms a lower arc, 
4.4 x 1.3 mm, with very obscure vertical lineations. 

The more slender jaw of A. craveni forms a fulvous, 
somewhat depressed arc, 4.5 x 0.7 mm, with ca 35 fairly 
uniformly distributed vertical riblets. 

Acknowledgements. Special thanks are given to the curators and 
their assistants of 28 museums, and to three private collectors, all 
individually referred to in the Acronyms - Institutional & Personal 
Collections. I am especially grateful to Drs A.C. van Bruggen 
(Rijksmuseum van Natuurlijke Historic Leiden), B. Verdcourt 
(Royal Botanic Gardens, Kew), and W.F. Sirgel (Stellenbosch 
University) for reviewing the manuscript and offering valuable 
suggestions; and to Dr P.K.. Tubbs, Executive Secretary of the 
International Commission on Zoological Nomenclature, for consulta- 
tions on the Code. I am also indebted to many others who have 
provided important assistance, among them: Donald B. Sayner, 
Charlotte Ernstein and Virginia Childs of the Scientific Illustration 
staff at the University of Arizona for photographs in Figures 27-42, 
45-46, 52-57; the Photography Service staff of the British Museum 
(Natural History) for photographs in Figures 23-26, 43-44, 47-51; 
David L. Bentley of the Electron Microscope Facility, Division of 
Biotechnology, and Professor Michael A. McClure, both of the 
University of Arizona, for assistance in the SEM photographs; 
Emilee M. Mead of the University of Arizona Teaching Center who 
drew the map and prepared the photographic layouts; Yolanda 
Baldonado Whigham of the U.A. Department of Ecology and 
Evolutionary Biology for word processing; and my wife Eleanor who 
has helped in countless ways. 



34 



A.R. MEAD 




Figs 58-63 Dorsal and dorso-right lateral views of radulae: 58, 59 Callistoplepa barriana (SMF, O. Boettger). 60, 61 C. shuttleworthi 
(SMNH no. 10). 62, 63 Leptocala mollkella (MRAC no. 795.638). 



NEW SUBFAMILY AND GENUS ACHATINIDAE 



35 





V 


\ 




- 


x** 1 




R^ 


6^, 


1 









Figs 64, 65 Bequaertina pintoi (BMNH no. 1953.8.15.562-564, mixed lot). 66, 67 B. graueri (MRAC no. 204.633). 68, 69 Achatina 
craveni (BMNH no. 1953.8.15.562-564, mixed lot). Bar scale = 1 fjun. 



36 



A.R. MEAD 



ACRONYMS - INSTITUTIONAL & 
PERSONAL COLLECTIONS 

AMNH New York: American Museum of Natural History (W.K. 

Emerson) 
ANSP Philadelphia: Academy of Natural Science (A.E. Bogan) 
BMNH London: British Museum (Natural History) (P. Mordan, 

F. Naggs) 
BV Kew, Royal Botanic Gardens: Bernard Verdcourt 

CMNH Pittsburg: Carnegie Museum of Natural History (J.E. 

Rawlins) 
FMNH Chicago: Field Museum of Natural History (A. Solem) 
GNM Goteborg: Naturhistoriska Museet (I. Levinsson, H.W. 

Walden) 
HM Newquay, Cornwall, England: Hazel Meredith 

IRSN Bruxelles: Institut Royal des Sciences Naturelles (J. van 

Goethem) 
LNK Karlsruhe: Landessammlungen fur Naturkunde (H-W. 

Mittmann) 
MCZ Harvard: Museum of Comparative Zoology (K.J. Boss) 
MNHN Paris: Museum National d'Histoire Naturelle (S. & A. 

Tillier) 
MRAC Tervuren: Musee Royal de l'Afrique Centrale (P.L.G. 

Benoit, F.A. Puylaert) 
NG Blantyre, Malawi: W. Noel Gray 

NHMB Bern: Naturhistorisches Museum (J.J. Oberling) 
NHMW Wien: Naturhistorisches Museum Wien (E. Wawra, O.E. 

Paget) 
NM Pietermaritzburg, South Africa: Natal Museum (R.N. Kil- 

burn) 
NMB Basel: Naturhistorisches Museum (C. Stocker- 

Unternahrer) 
NMW Cardiff: National Museum of Wales (A. Trew, P.G. 

Oliver) 
RMNH Leiden: Rijksmuseum van Natuurlijke Historie (A.C. van 

Bruggen, E.Gittenberger) 
RMS Edinburgh: The Royal Museum of Scotland (D. Heppell) 
SAM Cape Town: South African Museum (J. Pether) 
SMF Frankfurt: Forschungsinstititut Senckenberg Natur- 

Museum (R. Janssen) 
SMNH Stockholm: Naturhistoriska Riksmuseet (A. Waren, C. 

Holmquist) 
UHZI Hamburg: Universitat Zoologisches Institut und Museum 

(R. von Cosel) 
UMMZ Ann Arbor: University of Michigan Museum of Natural 

History (J.B. Burch) 
USNM Washington, D.C.: U.S. National Museum of Natural 

History (R. Hershler) 
UUZM Uppsala: Universitets Zoologiska Museum (A. Franzen, 

L. Wallin) 
ZMB Berlin: Museum fur Naturkunde der Humboldt- 

Universitat (R. Kilias) 
ZMUC K0benhavn: Zoologisk Museum - Universitet (J. Knud- 

sen, T. Schi0tte) 
ZSM Munchen: Zoologische Staatssammlung (R. Fechter) 



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Bull. not. Hist. Mus. Lond. (Zool.) 60(1): 39-104 



Issued 23 June 1994 



On Recent species of Spiraserpula 
Regenhardt, 1961, a serpulid polychaete 
genus hitherto known only from Cretaceous 
and Tertiary fossils 

T. GOTTFRIED PILLAI 

Marine Biological Services Division, Department of Zoology, The Natural History Museum, Cromwell 
Road, London SW7 5 BD 

HARRY A. TEN HOVE 

Universiteit van Amsterdam, Instituut voor Taxonomische Zoologie, Postbus 94766, 1090 GT 
Amsterdam, Nederland 

CONTENTS 



Synopsis 39 

Introduction 39 

Methods and materials 40 

Terminology 40 

Diagrammatic representations of internal tube structures 41 

Diagnosis of Spiraserpula Regenhardt , 1961 41 

Key to the Recent species of Spiraserpula Regenhardt, 1961 46 

Description of species 49 

Discussion 99 

Acknowledgements 103 

References 103 



Synopsis. A group of Recent serpulid species related to the genus Serpula Linnaeus, 1758, but differing from it in 
two important characters, is described in this paper. The first is a hitherto undescribed character, namely, the 
possession of internal tube structures which consist of longitudinal ridges and other structures, the form and 
arrangement of which, in combination with characters of the worms themselves, served to separate the various 
species. The second is that the thoracic membranes of the two sides in the worm do not unite ventrally at the end of 
the thorax to form a flap or apron as in Serpula. These characters are also common to 18 species, including three 
previously described ones. On the basis of the tube structures, these Recent species can be referred to the genus 
Spiraserpula Regenhardt, 1961, which was previously known only from fossils (Pillai, 1993). Scissiparity was 
observed in at least three of its species. A key to the known Recent species of Spiraserpula and a discussion on the 
systematics of the genus are included. 



INTRODUCTION 



In the course of a study of the serpulids currently referred to 
the genus Serpula Linnaeus, 1758, it was observed that the 
worms of larger species could frequently be extracted undam- 
aged from the anterior ends of their tubes with a pair of fine 
forceps, while they were invariably damaged in the process in 
certain small species as, for instance, in the well-known 
Mediterranean species Serpula massiliensis Zibrowius, 1968. 
In almost every collection of the latter, the worms which had 
been previously extracted from their tubes were incomplete 
posteriorly, and the ends of their longitudinal musculature 



provided evidence of their having been forcefully broken off 
from the rest of the abdomen. The cause of the difficulty in 
extracting complete worms was revealed by opening their 
tubes carefully from their anterior ends all the way to their 
posterior ends. It was found that the posterior end of the 
abdomen was retracted very tightly into the posterior coiled 
part of the tube and, quite surprisingly, against a longitudinal 
row of sharp serrations projecting from the inside of the tube. 
Examination of more material showed that these were consis- 
tent for 5. massiliensis. 

Study of similar material from various other geographical 
localities revealed the existence of species with other forms of 
internal tube structures. Evidently, these serve for anchorage 



)The Natural History Museum, 1994 



40 



T.G. PILLAI AND HA. TEN HOVE 



of the worm when withdrawn into the tube, and thereby, 
have an additional protective function. The form and 
arrangement of the internal tube structures, in combination 
with characters of the worms themselves, served to separate 
the various species. They are absent in Serpula Linnaeus, 
1758, and have not been described in any of the other known 
genera of Serpulidae. They differ from the transverse tabulae 
of certain serpulids, an account of which is given by Lom- 
merzheim (1979). Another important character common to 
the group is that, unlike in Serpula, the thoracic membranes 
of the two sides are not united posterior to the thorax to form 
a ventral flap or apron. 

In the search for a name for this group, the genera 
Pseudoserpula Straughan, 1967 and Protoserpula Uchida, 
1978, were considered, among others. The former was found 
to be invalid, and an account of the study which led to this 
conclusion is provided under Spiraserpula minuta, 
(Straughan, 1967), in this paper. It was not possible to 
examine the type specimen of Protoserpula to establish 
whether it has ITS or not. It is not in the National Science 
Museum, Tokyo, and other efforts to locate it were unsuc- 
cessful. 

H. Zibrowius of Station Marine d'Endoume, Marseille, 
who went through the manuscript of this paper, and the 
second author discussed the group with M. Jager of Rohrbach 
Zement, Dotternhausen, Germany, who re-examined the fos- 
sil serpulids studied by him (Jager, 1983), and other material, 
and found that some of them too possessed internal tube 
structures, although they had not been reported earlier. The 
collaboration which followed (pers. comm.) led to a study of 
likely Cretaceous and Tertiary serpulid genera and species 
(Pillai, 1993), which revealed that the group belongs to the 
genus Spiraserpula Regenhardt, 1961, previously known only 
from fossil species. Spiraserpula Regenhardt, 1961, has prior- 
ity over Protoserpula Uchida, 1978, even if the latter were to 
possess internal tube structures, henceforth referred to in the 
text as ITS {vide Jager 1993)). Zibrowius (1972) described a 
Recent spirorbid species belonging to the genus Neomicrorbis 
Rovereto, 1904, which was previously known only from 
Cretaceous and Tertiary fossils. 

In three of the Recent species of the genus Spiraserpula 
definite proof of asexual reproduction was found, in the form 
of branching tubes, corroborated by the presence of a parent 
with a schizont in one tube of Spiraserpula snellii sp. nov. 
Asexual reproduction had previously been reported for the 
genera Filograna Berkeley, 1835, Filogranula Langerhans, 
1884, Josephella Caullery & Mesnil, 1896, Salmacina Cla- 
parede, 1870 and Rhodopsis Bush, 1905 (ten Hove, 1979; 
Ben-Eliahu & ten Hove, 1989). Pillai (1993) reports the 
occurrence of tube branching in the fossil species Spiraserpula 
versipellis Regenhardt, 1961. It would not be surprising if it 
turns out that scissiparity takes place in most, if not all, 
species of the genus Spiraserpula, in view of their aggregated 
occurrence. 

Nineteen species of Spiraserpula, including the three 
known ones referred to above and an unnamed one, are 
described. They come from the Mediterranean, Madeira, 
Canary and Cape Verde Islands, Gulf of Mexico, the Carib- 
bean and Panama, the northern Red Sea, Mozambique, the 
eastern islands of Indonesia, Eastern Australia, Japan and 
New Caledonia. 



METHODS AND MATERIALS 



The tubes and their internal structures, as well as whole 
worms and parts were examined and drawn under a stereo 
microscope fitted with a drawing attachment. Measurements 
were taken with a pair of fine dividers against a scale having 
an accuracy of 0.5 mm, of total length of the tube when 
possible, external diameter of the tube, total length of the 
worm, width of the thorax just posterior to the pair of collar 
fascicles, length and diameter of operculum, length of the 
opercular peduncle, and length of the longest radiole and its 
pinnule-free tip when present. Radioles and thoracic seg- 
ments were counted on both sides, while the abdominal 
uncinal tori of one side were counted to determine the 
number of abdominal segments. The chaetae were mounted 
in polyvinyl lactophenol or aquamount and figured under the 
oil immersion lens of a high power microscope fitted with a 
drawing attachment. Measurements of chaetae were made 
with an eyepiece micrometer standardised with a stage 
micrometer. Scanning electron micrographs of chaetae of 
some of the species are also provided (Plates 1-5). 

The sources of material have been detailed under the 
respective descriptions as well as the acknowledgements. Full 
details of E. Atlantic stations surveyed by the 'Tydeman' 
Canary and Cape Verde Islands Expeditions of 1980, 1982 
and 1986 (CANCAP-IV, VI and VII), (e.g. CANCAP 
4.D14, 6.134) can be found in van der Land (1987); of E. 
Indonesian stations sampled during the Indonesian-Dutch 
Snellius II Expedition (e.g. Snellius II 4.051) in van der Land 
& Sukarno (1986). The following abbreviations have been 
used in the text: AM: Australian Museum, Sydney; AMNH: 
American Museum of Natural History, New York; BM(NH): 
British Museum (Natural History), London, presently. The 
Natural History Museum, London; FSBC I: Florida Depart- 
ment of Natural Resources, Invertebrate collection, St. 
Petersburg, Florida; HUJ: The Hebrew University, Jerusa- 
lem; MCZ: Museum of Comparative Zoology, Harvard; 
NNM: Nationaal Natuurhistorisch Museum, Leiden (for- 
merly Rijksmuseum van Natuurlijke Historie); NSMT: 
National Science Museum, Tokyo; QM: Queensland 
Museum, Brisbane; RMNH: Collection numbers of NNM; 
SME: Station Marine d'Endoume, Marseille (most material 
will be deposited later in the Musee Nationale d'Histoire 
Naturelle, Paris); USNM: United States National Museum of 
Natural History, Washington DC; V.Pol: Polychaete collec- 
tion numbers of ZMA; ZLU: Zoological Laboratory, 
Utrecht; ZMA: Zoologisch Museum, Instituut voor Taxo- 
nomische Zoologie, Amsterdam; ZMH; Zoologisches Insti- 
tut und Zoologisches Museum, Hamburg; ZMK: Zoologisk 
Museum, Kobenhavn. 



TERMINOLOGY 



The terminology used in this paper is explained in Figs 1 and 

2. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 

Hi 



41 







Plate 1 Scanning electron micrographs of fractured ends of tubes showing internal structures. A, C & D, Spiraserpula lineatuba (Straughan, 
1967). B, S. ypsilon sp. nov. 



DIAGRAMMATIC REPRESENTATIONS OF 
TUBES 



The various arrangements of ITS in the species described are 
diagrammatically represented in Fig. 3. 



DIAGNOSIS OF SPIRASERPULA Regenhardt, 
1961 

The original generic diagnosis of Spiraserpula was based only 
on the tube of its fossil type species, S. Spiraserpula Regen- 
hardt, 1961. Pillai (1993) provides an emended definition for 
fossil species based on characters of the tube. However, the 
recent species described here are distinguishable not only by 
characters of their tubes but also of the worms themselves, 



42 



T.G. PILLAI AND HA. TEN HOVE 




Plate 2 Scanning electron micrographs of chaetae. A-D, Spiraserpula massiliensis (Zibrowius, 1968): A, bayonet chaetae. B, thoracic uncini. 
C, anterior abdominal uncini. D, flat trumpet-shaped abdominal chaetae. E & F, 5. singularis sp. nov.: E, bayonet chaetae. F, abdominal 
uncini. 



and they have been taken into consideration in the following 
diagnosis: 

Tube with internal structures, usually towards its earlier 
formed, coiled, posterior portions. They consist of internal 
longitudinal ridges which vary in form and complexity in the 
different species; they may be dorsal, along the convex inner 



wall of the tube, and/or ventral, along the opposite side. They 
may be laminar, serrated or unserrated, or have other forms, 
and accessory lateral ridges or other structures may also be 
present. An umbilicus and peristomes may be present. There 
is usually an external granular overlay which bonds together 
coils of individual tubes, or those of other tubes to form 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



43 




Plate 3 Scanning electron micrographs of chaetae. A-D, Spirserpula nudicrista sp. nov.: A, bayonet chaeta. B, thoracic uncini. C, 
abdominal uncini. D, flat trumpet-shaped abdominal chaetae. E-G, S. lineatuba (Straughan, 1967): E, bayonet chaetae. F, abdominal 
uncini. G, flat trumpet-shaped abdominal chaetae. 



44 



T.G. PILLAI AND H.A. TEN HOVE 







Plate 4 Scanning electron micrographs of chaetae. A-D, Spiraserpula zibrowii sp. nov.: A, bayonet chaetae. B, thoracic uncini. C, 
abdominal uncini. D, flat trumpet-shaped abdominal chaetae. E & F, S. caribensis, sp. nov.: bayonet chaetae. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



45 





09KU 2.31KX 4.33H 8035 



B 




til 
ro 

h 




k 


(9 


Ni- 




\ 







Plate 5 Scanning electron micrographs of chaetae. A-C, Spiraserpula caribensis sp. nov.: A, abdominal uncini. B, flat trumpet-shaped 
abdominal chaetae. C-E, collar chaetae from Grenada material. F-H, 5. snellii sp. nov.: F, bayonet chaetae. G, abdominal uncini. H, flat 
trumpet-shaped abdominal chaetae. 



46 



T.G. PILLAI AND HA. TEN HOVE 



d.cx.w. 
dr. 




Fig. 1 Terminology. A, Tube: d.cx.w., dorsal convex wall; d.r., dorsal ridge; v.cv.w., ventral concave wall; v.r., ventral ridge. B, Radioles 
of both sides and operculum: o.p., opercular peduncle; p., pinnules; pf.t., pinnule-free tip; r., radius; rl., radiole; r.l.s., radioles of left 
side; r.o., rudimentary operculum; r.r.s., radioles of right side; t.r.l. triangular radial lobe; z.o., zygomorph operculum. C-G, Opercula: 
en., constriction between operculum and peduncle; i.r.g., inter-radial groove; ut.c, unthickened cuticle, bs.o., bell-shaped operculum; 
r.r.l., rounded radial lobe; t.t.c., thickened transparent cuticle. 



mutually bonded aggregations of a few to numerous individu- 
als. 

An operculum similar to that in Serpula, which is a 
modification of the second most dorsal radiole, is often 
present on one side with, correspondingly, a rudimentary 
operculum on the other. There may only be a rudimentary 
operculum on each side in certain species, while they may be 
present in juveniles and completely lost in older specimens in 
others. The shape of the fully developed operculum is charac- 
teristic for a particular species; it may be funnel-shaped, 
bell-shaped, zygomorphic or spherical. Its distal end may be 
concave or convex and usually bears radii which end as 
triangular or rounded lobes at the rim; but radii may also be 
lacking in some species. Its cuticle may be unthickened or 
thickened and transparent. 

The number of branchial radioles is usually small, rarely up 
to 14 pairs. Palps absent. A pair of prostomial ocellar clusters 
is usually present. The number of thoracic chaetal tufts may 
exceed the seven pairs commonly occurring in many genera 
of Serpulidae, including Serpula, and those of the two sides 
are more frequently asymmetrical than symmetrical. Up to 14 
have been counted on each side. Histological work is needed 
to ascertain the real extent of the segments, and their relation 
to numbers of chaetal tufts and uncinal rows, etc. The term 
'chaetiger' is, therefore, used here in the literal meaning of 
'hair bearer' and not as a synonym of segment. The thoracic 



membranes end anterior to the last thoracic chaetigers, also 
more frequently asymmetrically than symmetrically. Unlike 
in most species of Serpula sensu stricto, therefore, a post- 
thoracic ventral flap (apron) is absent. 

Collar fascicles bear chaetae of two kinds: bayonet 
chaetae and limbate chaetae, the blades of both of which 
are usually finely serrated. In the former, there are a few to 
several comparatively large teeth, located at the distal end 
of the shaft, separated from the bayonet-like blade by an 
unserrated area (unserrated notch). The range in the 
number of such teeth and the length of the unserrated 
notch varies in the different species. Limbate chaetae bear 
simple, more or less curved, blades. Thoracic and anterior 
abdominal uncini may bear teeth in a single row (saw- 
shaped), or are partly (saw- to rasp-shaped) or completely 
rasp-shaped. Abdominal chaetae bear distally flat trumpet- 
shaped ends, and are replaced by capillaries in the poste- 
rior segments. The distal ends of the abdominal chaetae of 
Serpula have been described as 'trumpet-shaped' in ser- 
pulid literature. We have discussed the inappropriateness 
of the comparison, as demonstrated by the scanning elec- 
tron micrographs and drawings of these chaetae presented 
in this paper, and our attention has also been drawn to this 
by Zibrowius (pers. comm.). In order not to create confu- 
sion, it was decided to retain 'flat trumpet-shaped', for the 
present. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



47 




Fig. 2 Terminology. A & B, Body of worm: a.t., abdominal tori; cap., capillary chaetae; col., collar; col.s., collar chaetae; d.l.g., dorsal 
longitudinal groove; t.s., thoracic chaetae; t.t., thoracic tori; v.l.g., ventral longitudinal groove. C-G, Bayonet-shaped collar chaetae (after 
ten Hove & Jacobs, 1984; all same magnification): act., accessory teeth; b.b., basal boss; f.t., few teeth; l.s.b.Jong serrated blade; 
m.l.s.h., moderately long serrated blade; s.s.b., short serrated blade; s.t., several teeth; u.n., unserrated notch; v.s.dl.b., very short 
dagger-like blade; v.s.t.b., very short tapered blade. H, Uncini, showing orientation in relation to the body of the worm: a., anterior; p., 
posterior. 



KEY TO THE KNOWN RECENT SPECIES OF 
SPIRASERPULA REGENHARDT, 1961 

(See Figure 3 for terminology of ITS) 

1. Tube with either dorsal or ventral internal longitudinal ridge 
only 2 

Tube with dorsal and ventral internal longitudinal ridges .... 7 

2. Tube with dorsal longitudinal ridge only (Fig. 3, B-E) 3 



Tube with ventral longitudinal ridge only, exceptionally with 
few isolated teeth (Fig. 3, F) 6 

Dorsal ridge unserrated, shaped like an inverted V (Fig. 3, B) 
S. singularis sp. nov. p. 62 

Dorsal ridge serrated (Fig. 3, C-E) 4 

Serrations of dorsal ridge deltoid (Fig. 3, C) 
S. deltoides sp. nov. p. 80 

Serrations of dorsal ridge not deltoid (Fig. 3, D & E) 5 



48 



T.G. PILLAI AND HA. TEN HOVE 

d.s. 




Fig. 3 Diagrammatic representations of ITS in the various species. A, Generalized drawing with all the main ITS. The orientation of the 
tube and terminology used are the same for all the diagrams. Ant., anterior direction, d, dorsal side, d.cx.w., dorsal convex wall.d.l.r., 
dorso-lateral ridge, d.r., dorsal ridge, v, ventral side.v.cv.w., ventral concave wall, v.r., ventral ridge. B, S. singularis sp.nov. C, 5. 
deltoides sp. nov.; d.s., deltoid serrations. D, S. massiliensis (Zibrowius, 1968). E, S. capeverdensis sp. nov.; k.c.d., knob-like calcareous 
deposits. F, 5. nudicrista sp. nov. & 5. snellii sp.nov.; v. v.r., variant form of ventral ridge. G, S. ypsilon sp. nov. & S. paraypsilon sp. nov. 
H, S. sumbensis sp. nov. I, S. iugoconvexa sp. nov. J, S. vasseuri sp. nov. K, S. plaiae sp. nov. L, S. caribensis sp. nov. & 5. lineatuba 
(Straughan, 1967). M, S. discifera sp.nov.; t.d., transparent discs. N, S. karpatensis sp. nov. & S. minuta (Straughan, 1967). O, S. zibrowii 
sp. nov. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



49 



5. Maximum number of radioles 6 pairs, abdominal segments 
about 50. Operculum may or may not be present. A shallow 
water species, down to about 60 m, rarely in deeper water 
S. massiliensis (Zibrowius, 1968) p. 49 

Maximum number of radioles 8 pairs, abdominal segments 
about 145. Operculum absent. A deep water species occurring 
at depths of about 75-200m S. capeverdensis sp. nov. p. 54 

6. Tube creamish white, trapezoidal in cross-section. Operculum 
absent. Maximum number of radioles 9 pairs, pinnule-free tips 
very long. Prostomial ocelli prominently seen through collar. 
Unserrated notch of bayonet chaetae very short 

S. nudicrista sp. nov. p. 76 

Tube mustard coloured, circular in cross-section. Operculum 
present. Maximum number of radioles 5 pairs, pinnule-free tips 
short. Prostomial ocelli not seen through collar. Unserrated 
notch of bayonet chaetae moderately long 
S. snellii sp. nov. p. 84 

7. Dorsal ridge serrated, ventral ridge Y-shaped (Fig. 3, G) ... 8 
Dorsal ridge unserrated, ventral ridge having other forms ... 9 

8. Thoracic uncini without lateral tubercles. Maximum number of 
radioles 7 pairs, abdominal segments more than 100, about 
130 S. ypsilon sp. nov. p. 56 

Thoracic uncini with lateral tubercles. Maximum number of 
radioles 11 pairs, abdominal segments less than 100, about 90 
S. paraypsilon sp. nov. p. 60 

9. Ventral ridge unserrated (Fig. 3, I) 10 

Ventral ridge serrated (Fig. 3, J-0) 11 

10. Tube white to faintly pinkish, circular in cross- section, external 
longitudinal ridges absent; dorsal and ventral internal longitudi- 
nal ridges pink, wedge-shaped in cross-section (Fig. 3, H). 
Operculum with up to about 21 triangular radial lobes, cuticle 
unthickened. Maximum number of radioles 5 pairs, abdominal 
segments below 100 (about 70); bayonet chaetae with several 
teeth on the basal boss S. sumbensis sp. nov. p. 82 

Tube bright rose, quadrilateral to trapezoidal in cross-section, a 
pair of external longitudinal ridges, and a faint median one in 
places; dorsal and ventral internal longitudinal ridges white, 
dorsal ridge T-shaped in cross-section, ventral ridge very small 
(Fig. 3, I), may or may not be present. Operculum with up to 
about 12 rounded radial lobes, cuticle unthickened. Maximum 
number of radioles 14 pairs, abdominal segments over 100 
(about 120); bayonet chaetae with two teeth on the basal boss 
S. iugoconvexa sp. nov. p. 82 

11. Bayonet chaetae with long blades and several teeth on the basal 
boss S. vasseuri sp. nov. p. 78 

Bayonet chaetae with short to moderately long blades and few 
(2-7) teeth on the basal boss 12 

12. Accessory lateral ridges present (Fig. 3, K & L) 13 

Accessory lateral ridges absent (Fig. 3, M-O) 15 

13. Accessory ridges dorso-lateral. Dorsal ridge wedge- to 
Y-shaped in cross-section (Fig. 3, K), tube white 

5. plaiae sp. nov. p. 67 

Accessory ridges lateral. Dorsal ridge a simple plate, at most 
wedge to faintly T-shaped in cross-section (Fig. 3, L), tube pink 
or with pink longitudinal bands 14 

14. Bayonet chaetae dagger-shaped, with short blunt blades. Oper- 
culum absent. Maximum number of radioles 6 pairs 

S. caribensis sp. nov. p. 68 



Bayonet chaetae with moderately long blades and tapering tips. 
Operculum present. Maximum number of radioles 5 pairs; 
S. lineatuba (Straughan, 1967) p. 91 

15. Tube with transparent discs attached to the wall externally and 
internally (Fig. 3, M) S. discifera sp. nov. p. 94 

Tube without such discs (Fig. 3, N & O) 16 

16. Operculum present S. karpatensis sp. nov. p. 64 

Operculum absent 17 

17. Bayonet chaetae with 5-7 teeth on basal boss; maximum 
number of radioles 4 pairs, abdominal segments about 55 

S. zibrowii sp. nov. p. 67 

Bayonet chaetae with 3-4 (rarely 5) teeth on basal boss; 
maximum number of radioles 6 pairs, abdominal segments 
about 80 S. minuta sp. nov. p94 

Although the ITS are very distinctive, the states of the 
characters need to be used with caution. In two species S. 
lineatuba (Straughan, 1967) and S. caribensis sp. nov., for 
example, 25^10 tube fragments had to be examined before 
the full extent of the development of dorsal and accessory 
ridges could be established; the latter are missing in most 
cross-sections. The shape of the distinctive inverted V, as in 
the dorsal ITS of 5. singularis sp. nov. is only found in a small 
section in the earlier formed part of the tube; elsewhere, the 
ridge is a smooth plate only. Along this ridge, the rounded 
edge gradually becomes indented, gutter-shaped, and finally 
widening to form a V. This would apply to certain other 
characters as well. In 5. massiliensis (Zibrowius, 1968) part of 
the sample from Marseille was operculate and part had 
rudimentary opercula only. However, all the specimens from 
a large sample from Portman had rudimentary opercula only. 
It may thus be expected that species which, on the basis of 
relatively few specimens, have been described as non- 
operculate, may turn out to be operculate when more mate- 
rial becomes available. As another example, two samples 
from Indonesia and Lizard Island (Queensland) initially 
appeared to belong to two distinct species, on the basis of 
differences in six character states. Additional material, how- 
ever, yielded specimens with a full range of intermediate 
states, showing that they belong to one and the same species. 



DESCRIPTION OF SPECIES 

Spiraserpula massiliensis (Zibrowius, 1968) 
(Figs. 4, A-O; 3, D; PI. 2, A-D) 

SYNONYMY. Serpula massiliensis Zibrowius, 1968: 102-105, 
Pl.l, figs.24-37; P1.14, fig.d. 

Serpula massiliensis: Bianchi, 1981 : pp. 51-52, fig. 16. Serpula 
massiliensis: ten Hove & Aarts, 1986: 35 [not the tropical E. 
Atlantic record, see S. ypsilon]. 

Material examined. Unless otherwise mentioned, the 
material was collected and/or determined by Zibrowius. 
Mediterranean: 

France: Marseille: 1. Anse des Cuivres; below SME, over- 
hang 6m, 21.vii.1987 (10 out of several specimens, BMNH 
ZB 1989, 43-53). 2. tie Plane; submarine cave, 6m, legit G. 
Harmelin, vi.1987 (4 specimens, SME). 3. lie Plane; 1987 (5 
out of several specimens, BMNH 1989 101-150).4. lie Plane; 



50 



T.G. PILLAI AND HA. TEN HOVE 




Fig. 4 Spiraserpula massiliensis (Zibrowius, 1968). A-O, From Marseille, Anse des Cuivres, BM(NH). ZB1989. 43-53. A, Aggregation of 
tubes with fractured ends showing the serrated dorsal ridge along the convex inner wall, and granular overlay. B-C, Erect parts of tube 
with four-lobed peristome. D, Anterior part of operculate worm. E, Same specimen showing end of thoracic membrane. F, Thorax with 
pair of prostomial ocellar clusters, also enlarged. G-J, Four bayonet chaetae from the same fascicle. K, Row of thoracic uncini. L, Anterior 
abdominal uncini. M & N, Middle abdominal uncini. O, Posterior abdominal uncini. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



51 



submarine cave, 6m, legit G. Harmelin, vii.1971 (7 speci- 
mens, ZMA V. Pol. 3159). 5. Friocil Harbour (5 out of 
several specimens, BMNH ZB 1989 54-100). 6. Grand Con- 
glu; 1987 (5 out of several specimens, BMNH ZB 1989 
151-200). 7. Martigues; ca. 50km W of Marseille, Ponteau 
Electric Plant, under stones, 1.3m, 5. iv. 1977 (2 out of several 
specimens, SME). 8. La Ciotat; ca 30km E. of Marseille, Bee 
de l'Aigle, on concretions of sand, 40m, iii.1970 (3 speci- 
mens, SME). 9. Canyon de la Cassidaigne; about 20km E of 
Marseille, off Cassis, from 170-270m by dredging, 15. vi. 1974 
(tubes, 1 specimen, SME). 

NW Corsica: 10. Revellata, 15m, calcareous algal masses, 
8.iv.l978 (2 specimens, SME). 

Italy: 11. S coast of Sorrento peninsula, 'Grotto Zaffiro', 
10m, 29.V.1974 (3 specimens, SME). 12. Bari; 10m, cave, 
legit T.M.Griessinger, 8.vii.l968 (5 specimens, SME). 
Greece: 13. Gulf of Corinth, Aspra Spitia, 5m, 26. ix. 1977 (3 
specimens, SME). 

Malta: 14. Oxford University Underwater Exploration Group 
1965, scrapings from roof of cave, det. Pillai (2 specimens, 
BMNH ZB1989 32-36). 

Tunisia: 15. Tabarka; algal concretions, 31-36m, 24. iv. 1969 
(1 specimen, SME). 16. Zembra Islands, concretions, 35m, 
30. iv. 1969 (few tubes with portions of worms, SME). 17. 
'Dauphin' Stn.24, 35°12'N 11°25'E, 73m, on Area, legit 
Bane, Medit. Mar. Sorting Center, 28.viii.1967 (1 specimen, 
SME). 18. Gulf of Gabes; 'Calypso', 34°05'N 10°48'E, 23m, 
muddy sand with Caulerpa meadow, on shelly material, 
20.iv.1965 (1 specimen, SME1887). 19. Gulf of Gabes; 
'Calypso', 34°13'N 10°31.9'E, 31m, Caulerpa meadow, 
27.iv.1965 (1 specimen, SME 1910). 

SE Spain: 20. Cabo de Palos; ca 50km E of Cartagena, 6m, 
legit A. Ramos, 4.iv. 1982 (3 specimens, SME). 21. Portman; 
20km E of Cartagena, small overhang, 0.5-1.0m, on rock 
covered by dark brown sediment, the latter retained on the 
tube surfaces by oil pollution, 5.iv.l984 (20 out of several 
specimens, SME). 

Portugal: 22. From a submarine cave near Sagres, Algarve, 
legit H. Zibrowius Sept. 1986 (BM(NH) 1992. 181-255). 
NE Atlantic: 23. Gorringe Bank; 'Meteor' M9c, Stn.95, AT 
29, 36°29.9'N 11°33.0"W, 150-430m, 24. vi. 1967 (some empty 
tubes, SME). 24. Madeira Archipelago; Jean Charcot, 
Stn.42, SW of Porto Santo, approx. 33°0.4'N 16°24.5'W, 
125-145m, 17.vii. 1966 (empty tubes, information pers. 
comm. H. Zibrowius). 25. Canary Islands; W coast of Palma, 
Tijarafe, 28°42'N 17°58'W, 20m, CANCAP 4.D14, det. M. 
Aarts (5 out of several specimens, RMNH 18465, ZMA 
V.Pol. 3739, BM(NH) 175-180). 26. NW Africa; off Point 
Elbow, ex Spanish Sahara, 'Tenace' D16, Stn. 23, 24°13'N 
16°17'W, 50-60m, legit Marche-Marchard, 13. iv. 1967 (4 
specimens, SME). 

Type locality. Marseille (France). 

Description. In order to follow the variations within the 
genus Spiraserpula it was considered useful to have as com- 
plete a description as possible of one member of the group, S. 
massiliensis was selected because of the large amount of 
material available from various sources. The following 
updated species description is based on the original account 
as well as additional data obtained from a study of the above 
collections, which include much of Zibrowius' original mate- 
rial. 

According to Zibrowius (1968), the tubes are white, circu- 



lar in cross-section and, although difficult to measure because 
of their coiling, may attain a length of 50.0 mm for a diameter 
of about 0.5-1.0 mm. Their coiling is highly irregular and the 
direction may reverse. Sometimes many tubes are joined 
together, coiling in the same direction. They are relatively 
thick, except in their erect portions which are cylindrical. At 
intervals along the latter, there may be one to a few out- 
wardly directed expansions, generally referred to as peris- 
tomes. They are sometimes in the form of four, somewhat 
symmetrically placed lobes. In dense populations, the erect 
tubes may form a kind of uniform meadow in submarine 
caves of the Mediterranean (Zibrowius, pers. comm.). The 
surface of the tube is covered by faint granulations which, 
very rarely, may form short longitudinal ridges. 

It is difficult to remove the worms from their tubes. When 
removed, quite a number of specimens lacked their radiolar 
crowns. An operculum may be present or absent. When 
absent there is a rudimentary operculum on each side. When 
present, the operculum is small, and its diameter does not 
correspond with that of the tube. Its distal end is flattened to 
slightly funnel-shaped, bearing 13 to 23 obtuse lobes. The 
peduncle is more slender than the pinnulate radioles, and its 
attachment to the operculum is central and constricted. The 
corresponding radiole of the opposite side is reduced to a 
filamentous rudimentary operculum, about one-third the 
length of the radioles, and lacks pinnules. 

The collar consists of a large ventro-median lobe and a 
smaller one on either side of it, all of which are rounded. The 
thoracic membranes are broad and well developed up to 
about the fourth thoracic chaetiger, and reduced posteriorly. 
They are not united to form a post-thoracic ventral flap or 
apron. The number of thoracic segments may exceed the 
usual seven found in many other species of Serpula. Each 
collar fascicle generally possesses four bayonet-shaped 
chaetae and a similar number of simple bladed chaetae. Each 
bayonet chaeta has a striated blade distally, and several teeth 
on the basal boss. Thoracic uncini bear 3-5 teeth. Anterior 
abdominal bundles consist of 2-3 flat trumpet shaped 
chaetae. Uncini possess 2-5 teeth in a single row. The 
posterior abdominal segments bear long capillary chaetae, 
and rasp-shaped uncini with several rows of teeth. 

Additional data obtained during the present study are as 
follows: 

TUBES: White to faintly creamish, and may occur in closely 
intertwined masses of a few to several individuals (Fig. 4, A); 
sometimes solitary. Except for their free erect portions, they 
are mutually bonded to various extents, particularly at their 
bases. Their 'granular overlay' is shown in Fig. 4 A, and the 
four-lobed peristomes in Fig. 4, B & C. 

An important character which had not been reported 
relates to the tube, which bears ITS. In its first formed 
portion, which is normally coiled, there is a serrated longitu- 
dinal ridge. Careful removal of numerous specimens from 
their tubes has shown that this serrated ridge is always on the 
convex side of the coils, as also found in masses consisting of 
several individuals (Fig. 4, A). The orientation of the worm 
within the tube is such that the posterior dorsal part of its 
abdomen is always applied to this serrated 'dorsal ridge' 
(Fig. 3, D). This, in addition to its tight coiling, accounts for 
the difficulties encountered in extracting complete worms 
from their tubes by Zibrowius (1968) and in the present 
study. 

The numerous specimens in the collection from Portman 
show an apparent exception in lacking ITS. However, confir- 



52 



T.G. PILLAI AND HA. TEN HOVE 




ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



53 



mation that the specimens belong to this species came from a 
very small number of old and empty tubes at the bottom of 
the aggregations having the characteristic serrated internal 
dorsal ridge. The population had been subjected to oil 
pollution, as evidenced by a thick deposit of it which covered 
the tubes externally, and even lined them thinly internally. 
worms: The longest operculate specimen (from lie Plane, 
Marseille, BMNH ZB 1989 101-150) is 19.0 mm long; its 
thoracic width 1.1 mm. It has 5 pairs of radioles, in addition 
to an operculum on one side and a rudimentary operculum on 
the other. It also has the longest operculum and peduncle, 
3.5 mm; its operculum is 1.0 mm long, 1.2 mm in diameter, 
bell-shaped, and with 11 rounded marginal lobes. The abdo- 
men is 13.5 mm long and has 51 segments, the last four with 
capillaries. The longest non-operculate specimen, from the 
same locality, has a total length of 20.0 mm. Its abdomen is 
15.5 mm long and has 43 segments, the last ten with capillar- 
ies. The two specimens indicate variations in abdominal 
length due to extent of contraction during fixation, and that 
the number of segments in the abdomen may not be always 
proportional to its length. In the other specimens studied, the 
length of the operculum together with the stalk ranges from 
1.0-2.0 mm; the operculum from 0.5-0.6 mm in length and 
0.3-1.5 mm in diameter. 

The shape of the operculum varies from an elongate funnel 
in the majority of cases to a narrow bell in the others (Fig. 4, 
D, E).The opercular radii end in 10-16 rounded marginal 
lobes. The width of the peduncle, just before the constriction, 
is 3/5ths to 4/5ths that of the proximal end of the operculum. 
The number of radioles observed is 4-6 pairs, and their 
pinnule-free tips vary in length from l/5th to2/3rd the total 
length of the radiole (i.e. radiole plus pinnule-free filament). 
Zibrowius (1968) reported a higher maximum number of up 
to 23 marginal lobes on the operculum. This high number, in 
one specimen, is apparently not representative of the species 
(Zibrowius, pers. comm.). 

A total of 67 specimens from different localities (including 
21 from the abnormal Portman material, vide below) were 
examined for various characters. All the Portman specimens 
possess only rudimentary opercula. Out of 46 specimens from 
the other collections 25 possess an operculum, 12 lack one, 
and the rest are indeterminate since they lack radiolar 
crowns. The majority of specimens from normal populations, 
therefore, possess an operculum. 

Another character, hitherto not reported, is the presence 
of two light red to reddish-brown clusters of prostomial ocelli 
(Fig. 4, F). They can be seen when a worm with its radioles 
removed is viewed from the anterior end, or through the 
collar in mounted specimens. Each ocellus consists of a 
pigmented cup-shaped part, and a transparent anteriorly or 
antero-laterally directed lens-shaped part within it (Fig. 4, F). 
Thoracic glands, as found in other species of the genus, are 
absent. 

The numbers of thoracic chaetal tufts, 6 to 9 on each side, 
may be symmetrical or asymmetrical. The condition in 26 



Table 1 Spiraserpula massiliensis (Zibrowius). Number of thoracic 
chaetal tufts on each side. 



No. of individuals (n=26) 
No. of thoracic chaetal tufts 



1 

6/7 



4 

7/7 



7 
7/8 



11 

8/8 



3 
8/9 



Table 2 Spiraserpula massiliensis (Zibrowius). Extent of the 
thoracic membranes of the two sides. 



No. of individuals (n=21) 
Thoracic membrane ends 



1 


2 


10 


2 


3 


3 


4/5 


4/6 


5/5 


5/6 


6/6 


6/7 



specimens is summarized in Table 1. Likewise, the thoracic 
membranes may end symmetrically or asymmetrically, but 
always anterior to the last thoracic chaetiger; an apron is, 
therefore, absent (Fig. 4, E). The condition in 21 specimens 
is given in Table 2. 

Collar fascicles bear 3 to 5 bayonet chaetae each. Bayonet 
chaetae consist of a long serrated blade, an unserrated notch 
of moderate length, and a basal boss with several teeth of 
variable size (Fig. 3, G-J; PL 2, A). Thoracic and anterior 
abdominal uncini usually have 4 or 5 teeth arranged in a 
single row (edge saw-shaped), (Fig. 4, K, L; P1.2, B & C). In 
the intermediate abdominal region, the edge of each uncinus 
is saw-shaped anteriorly whereas several teeth are placed side 
by side (edge rasp-shaped) posteriorly. The number of teeth 
in a single row decreases and the rasp-shaped posterior 
portion increases as the posterior end of the abdomen is 
reached (Fig. 4, M-O). Although the posterior abdominal 
uncini are rasp-shaped, they have a single large tooth anteri- 
orly (Fig. 4, O). 

Live material. (Vide Zibrowius, 1968) 

Habitat and distribution. This species is commonly 
found in submarine caves and at depths accessible by diving 
(Zibrowius, 1968). The original description mentioned a 
depth of 10-22m, but subsequently the species was found to 
occur in shallower and deeper water (see list of material 
examined). Deeper water collections came from depths of 
31-36m (Tunisia) and 58-60m (off Point Elbow, Western 
Sahara), the latter consisting of operculate and non- 
operculate specimens. Empty tubes of this and other serpulid 
species typical of shallow water have been obtained along the 
the steep slope of the Gorringe Bank at 150-430m. This 
occurrence may be due to slumping from shallower depths 
(Zibrowius, pers. comm.). The Madeira Archipelago mate- 
rial (125-145m) also consisted of dead material. The empty 
tubes and single specimen from Canyon de la Cassidaigne 
(170-200m) is also exceptional. In general, therefore, the 
species commonly occurs in depths to about 60m, rarely down 
to about 200m. 

S. massiliensis is common in the Mediterranean (Greece, 



Fig. 5 Spiraserpula capeverdensis sp. nov. A-P, From type locality (SW of Sao Vicente), CANCAP 6.148 & 6.146; A-L, From 6.148, M-P, 
from 6.146. A, Opened tube showing serrated internal dorsal ridge along the convex wall of coiled part; and granular overlay in places. B, 
Coiled part of tube with fine transverse growth wrinkles externally, and its fractured end showing dorsal ridge on convex inner wall, and 
two ventro-lateral longitudinal rows of smooth tubercles on opposite wall. C & D, Cross-section of two tubes, both with serrated internal 
dorsal ridge, and one with ventro-lateral rows of tubercles. E-G, Three views of same worm showing rudimentary opercula (F), condition 
of the thoracic membranes (G), and dorsal longitudinal groove (E & G), and ventral abdominal groove posteriorly (E). H, Anterior end of 
younger specimen. I-L, Bayonet collar chaetae, all from same fascicle. M, Thoracic uncini. N, Anterior abdominal uncini. O, Uncini from 
transitional region of abdomen. P, Posterior abdominal uncini. 



54 

Italy, France, Spain, Malta, and Tunisia). In the eastern 
Atlantic it is abundant in submarine caves, and has been 
recognized on Gorringe Bank, the Madeira Archipelago, 
Portugal and the coast of Sahara. 

S. massiliensis has been erroneously reported from the Red 
Sea (Amoureux et ai, 1978). Examination of the specimens 
(HUJ) showed that their tubes lack ITS and they do not, 
therefore, belong to the genus Spiraserpula. 

Spiraserpula capeverdensis sp. nov. 
(Figs.5, A-P; 3, E) 

Material examined. 

Cape Verde Islands: All CANCAP stations. Off Sao Vicente: 

1. 6.134; 110-120m, (2 PARATYPES and some empty tubes, 
RMNH 18197). 2. 6.135; 110-150m, (1 PARATYPE, 
BM(NH) 1992.8). 3. 6.137; 75-90m, (1 PARATYPE, 
BM(NH) 1992.9). 4. 6.146; 75m, (1 specimen, BMNH). 5. 
6.148; 100-200m, (HOLOTYPE, 2 PARATYPES & 3 empty 
tubes (residual material) ZMA VPol. 3651). 6. 6.166; 
78-85m, (1 PARATYPE, USNM 130995). Off Razo: 7 
.7.117, 100-120m, (some empty tubes, RMNH 18198). 8. 
7.123; 120m, (5 specimens, RMNH 18199, ZMA V.Pol.3733. 
Scuba diving station : Boa Vista: 9. 7.D06; down to 12m, (3 
questionable specimens, ZMA V.Pol. 3871). 

Type locality. Cape Verde Islands, Sao Vicente. 

Description. 

TUBES: White, nearly circular in cross-section, and occurring 
in aggregations of a few individuals, occasionally solitary. 
They are closely coiled amongst or upon themselves (Fig. 5, 
A), and mutually bonded by a granular overlay. Erect 
portions, when present, are very short, hardly rising above 
the rest of the tube, and may end in four lobes. Faint growth 
rings are sometimes present (Fig. 5, B), and anterior 
uncoiled portions may sometimes show a few transverse 
thickenings, representing peristomes. In their first formed 
parts, they possess an internal serrated dorsal ridge (Fig. 5, 
A, D) and, often, a short ventro-lateral longitudinal row of 
small smooth knob-shaped processes on each side (Figs.5, B, 
C; 3, E). A mid-dorsal longitudinal groove in the posterior 
part of the abdomen (Fig. 5, E, G) is applied to the serrated 
dorsal ridge when the worm is withdrawn into the tube. The 
maximum external diameter of the tube varies from 0.6 mm 
in a juvenile to 1.4 mm in older specimens. 
worms: (Fig. 5, E-H). An operculum is absent in all the 
specimens examined. Instead, a filamentous rudimentary 
operculum is present on each side. The number of radioles in 
the larger specimens is often 7 or 8 per side, 4 in the smallest. 
They are about 2.0 mm long in the larger specimens, and 
have transverse specks at intervals. Their pinnule-free tips 
are slender, 1/5 to 1/6 the total length of the radioles. Two 



T.G. PILLAI AND HA. TEN HOVE 
Table 3 S. capeverdensis sp. nov. Measurements and meristic data. 



Stn. No. 


TL 


Width 


No. 


of 


Length 


No. of 


Caps. 




(mm) 


of 


radiol. 


of 


abdom. 


on 






thorax 






abdom. 


segs. 








(mm) 






(mm) 






6.137 


25.9 


0.5 


8/8 




22.7 


138 


27 


6.148 


20.6 


0.5 


7/8 




17.5 


145 


- 


6.148 


7.1 


0.5 


7/8 




6.9 


96 


12 


7.123 


2.9 


0.3 


4/4 




2.6 


49 


- 


7.123 


2.4 


0.3 


4/4 




2.2 


29 


9 



Table 4 S. capeverdensis sp. nov. Numbers of thoracic chaetal tufts 
and extent of thoracic membranes. 



No. examined (n = 12) 

No. of thoracic chaetal tufts 

No. examined (n=9) 
Thoracic membrane ends 



2 12 14 2 

9/8 8/8 8/7 8/5 7/7 7/6 

14 2 2 

6/6 5/5 5/? 4/4 



clusters of reddish to reddish-brown prostomial ocelli are 
present. 

Measurements and other data from the two longest and 
three juvenile worms are presented in Table 3. The numbers 
of thoracic chaetal tufts and the extent of the thoracic 
membranes on the two sides is variable, as shown in Table 4. 

The thoracic membranes do not extend to the last thoracic 
chaetigers (Fig. 5, F), and apparently end symmetrically, but 
further study of additional material is necessary for confirma- 
tion of the latter. Ventral thoracic glands are absent. 

Each collar fascicle bears up to about 5 bayonet chaetae 
(Fig. 5, I-L). They have a long serrated blade, a short 
unserrated notch and several moderately large teeth on the 
basal boss. Thoracic uncini (Fig. 5, M) usually have 4 teeth in 
a single row. Anterior abdominal uncini are similar, with 4-6 
teeth (Fig. 5, N). The posterior abdominal uncini are rasp- 
shaped, except for the single anterior tooth (Fig. 5, P). There 
is a transition (Fig. 5, O) between the condition found in the 
anterior and posterior abdominal uncini. 

The differences between S. capeverdensis sp. nov. and S. 
massiliensis are as follows: The former has only rudimentary 
opercula, and higher maximum numbers of radioles (8 pairs) 
and abdominal segments (145). Its tubes do not form tall 
erect portions, and usually possess two ventro-lateral rows of 
knob-shaped tubercles internally, in addition to the serrated 
dorsal ridge. In 5. massiliensis, however, an operculum may 
or may not be present, the maximum number of radioles is 6 
per side, and of abdominal segments observed 51. There is 
also strong indication of an ecological difference (see below). 



Fig. 6 Spiraserpula ypsilon sp. nov. From type locality material (SW coast of Island of Brava), CANCAP 6.D03. A, Aggregation of 
fractured tubes showing ITS, consisting of a Y-shaped ventral ridge along the concave wall and a serrated dorsal ridge along the convex 
wall. An oblique section (bottom right) shows the tapering anterior end of the ventral ridge. B, Erect portion of tube showing four-lobed 
peristome. C-E, Different views of complete worm showing showing rudimentary opercula, pinnule-free tips of the radioles (E), and dorsal 
and ventral longitudinal abdominal grooves. F-H, Same anterior end showing four-lobed collar (H), thoracic chaetigers and membrane (F), 
and ventral longitudinal groove. I-K, Three views of larger specimen, showing dorsal and ventral longitudinal abdominal grooves. Note the 
longitudinal cord-shaped structure within the ventral groove of the abdomen which fits into the gutter-shaped part of the Y-shaped ventral 
ridge of the tube. L, Anterior portion of of worm accidentally fixed outside its tube, showing filamentous rudimentary opercula and 
thoracic membranes. M-O, Three views of anterior part of another worm fixed outside its tube. Its thorax is considerably wider than those 
of specimens fixed within their tubes, and the longitudinal grooves may be stretched and shallow (N). 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



55 




56 



T.G. PILLAI AND H.A. TEN HOVE 



Remarks. Although the few damaged specimens from Stn. 
7.D06 are very similar to 5. capeverdensis in most respects, 
only 2 tube parts (out of 10 recognizable fragments) showed a 
serrated dorsal ridge and possibly latero-ventral knobs. As 
opposed to all material of S. capeverdensis studied so far, a 
specimen in a tube without visible internal structures showed 
a bell-shaped operculum with 13 radii. Therefore the identifi- 
cation of this lot is left at ? capeverdensis. 

Live material. No records. 

Etymology. Named after the type locality. 

Habitat and distribution. S. capeverdensis sp. nov. is 
known only from the Cape Verde Islands, mainly from 
depths of 75-200m where the bottom consists of various 
combinations of coarse sand, shell gravel, calcareous stones, 
calcareous nodules, calcareous algae and sponges, on which it 
occurs among the epifauna. 

Scuba diving to 15-20m, in a total of 28 different stations 
off the Cape Verde Islands during CANCAP-VI and CAN- 
CAP-VII, did not yield this species from the shallower 
coastal waters. However, the dives yielded a different spe- 
cies, S. ypsilon sp. nov., from these depths which, in the 
Mediterranean, are typical for S. massiliensis. 

Spiraserpula ypsilon sp. nov. 

Figs. 6, A-K; 7, A-T; 34, G; PI. 1, B) 

Synonymy. Serpula massiliensis: ten Hove & Aarts, 1986: 
35 (tropical E. Atlantic record only). 

Material examined. 

Cape Verde Islands: CANCAP stations. Scuba diving sta- 
tions: 1. 6.D01; S coast of Sao Tiago, SE of Porto Praia, 15m 
(1 specimen, RMNH 18177). 2. 6.D02; S coast of Sao Tiago, 
Baia de Santa Clara, 20m, caves in rock (2 out of several 
specimens; RMNH 18187; BM(NH) 1992.85-115; FSBC I 
39197 (1); AM W 20339 (1); NSMT (1)). 3. 6.D03; SW coast 
of Brava, Porto dos Ferreiros, 15m (30 specimens: HOLO- 
TYPE & 5 PARATYPES, RMNH 18176. Other 
PARATYPES: ZMA V. Pol. 3650 (10); USNM 130993 (6) 
and BM(NH) 1992.73-82 (10)). 4. 6.D06; SW coast of Sao 
Nicolau, Baia do Tarrafal, 15m (4 specimens, RMNH 18188). 
5. 6.D10; S. coast of Sao Vicente, 15m (5 out of several 
specimens, RMNH 18189). Coastal stations: 6. 6.K13; SW 
coast of Una Razo, (14 out of several specimens, RMNH 
18190, ZMA V.Pol.3726, USNM). 7. 6.K15; SW coast of Una 
de Santa Luzia, (1 out of several specimens; bulk RMNH 
18191; clusters of 10-15 tubes each BM(NH), 1992.116-120; 
ZMA V.Pol.3727; HUJ; Dr M. Jager). 8. 6.K21; NE coast of 
Sao Vicente, Baia das Gatas, (3 specimens, RMNH 18192). 
Scuba diving stations: 9. 7.D03; Cima, SE coast, (1 specimen, 
RMNH 18193). 10. 7.D05; Maio, SW coast of Ponta Preta (2 
out of few specimens, RMNH 18194). 11. 7.D06; Boa Vista, 
Ilheu de Sal Rei, 12m (1 out of few specimens, ZMA 
V.Pol.3728). 12. 7.D10; Razo, S coast, 20m (1 out of few 
specimens RMNH 18195; BM(NH) 1992. 121-131; ZMH). 
Dredging station:13. 6.148: off Sao Vicente, 100-200m (1 
empty eroded tube; RMNH 18196). Tropical Western Atlan- 
tic, Gulf of Mexico: 14. Florida, Stn. EJ66-460, 26°24'N 



82°28'W, 18m, 6.xii.l966, 'Hourglass' Stn J, (20 out of 
several specimens, FSBC I, ZMA V.Pol. 3729, BM(NH) 
1992. 132-147). 15. Florida: Stn. EJ 67-76, 27°37'N 83°28'W, 
39m, 2. hi. 1967, 'Hourglass' Stn.C, (few specimens, FSBC I, 
ZMA V.Pol.3730. 16. Florida: Stn. EJ67-328, 27°37N 
83°07W, 18m, ll.ix.1967, 'Hourglass' Stn. B (4 out of several 
specimens, ZMA V.Pol. 3731). Caribbean: 17. Aruba: Andi- 
curi, cape W of beach, windward side, rockpool, exuberant 
coral growth, strong wave action, 0.5m, legit H.A. ten Hove, 
28.viii.1970, Stn.2034B (together with 5. caribensis sp. nov.; 
ZMA V.Pol. 3732). 18. Colombia: Santa Marta area, Cabo 
and Ojo del Aguja, 8-27m, legit J. W. Dulfer and M. J. C. 
Rozenmeyer 1986, ident. as 5. massiliensis (1 damaged 
specimen, tube; ZMA V.Pol. 3778). Bermuda: 19. Stn. 14, 
legit Reed, with a note by Zibrowius in 1970 indicating that it 
is a new species (3 specimens, USNM 43244). 

Type locality. Cape Verde Islands, Brava. 

Description. 

tubes: Faintly pinkish, often with a more pronounced shade 
of light pink or light mauve in the granular overlay towards 
their anterior ends. They normally occur in mutually bonded 
highly coiled aggregations, occasionally also singly, adjacent 
to the aggregations. The granular overlay is fine, somewhat 
translucent and nearly uniform (partly shown in the top left 
portion of Fig. 6, A). External longitudinal ridges are nor- 
mally absent, but up to three may be faintly developed on the 
less coiled tubes of solitary specimens. Their anterior por- 
tions are generally attached, often with their lateral borders 
extending somewhat over the substratum. Occasionally, they 
possess erect ends which attain an external diameter of up to 
1.25mm, and may bear a few peristomes which are usually 
four-lobed and outwardly directed (Fig. 6, B). 

The ITS are more complicated than those of the other 
known species of the genus, with the exception of the closely 
related species S. paraypsilon sp. nov. As seen in carefully 
opened tubes or through their fractured ends in an aggrega- 
tion (Fig. 6, A), they consist of a serrated dorsal ridge, and a 
thin, very fragile, Y-shaped ventral ridge (Fig. 3, G; Pl.l, B). 
The gutter-shaped part and the stem of the latter gradually 
decrease anteriorly until they are represented only by a 
simple ventral ridge, which itself decreases in height and 
gradually disappears (Fig. 6, A, bottom right corner). These 
ridges commence in the first formed portions of the tube, but 
usually extend more anteriorly than in most of the other 
species of the genus. The inner translucent layer of the tube is 
faintly pinkish, as is the Y-shaped ventral ridge. In addition, 
the latter may possess one or two thin dark pink longitudinal 
stripes on the outside of the gutter-shaped part, one along the 
top and the other along the bottom. 

worms: Some measurements and counts are provided in 
Table 5-8. 

The worms attain a total length of about 27.5mm, a 
thoracic width of 0.6mm, a maximum of about 131 abdominal 
segments, with capillaries on the last 10 segments or so. The 
maximum number of radioles is 8 pairs. The two specimens 
with 4 pairs of radioles are juveniles. The pinnule-free tips 
are short to moderately long, up to about 1/5 the entire length 



Fig. 7 Spiraserpula ypsilon sp. nov. A-0, From type locality, CANCAP 6.D03. P-T, From Florida Stn. EJ 66-460. A-E, Bayonet chaetae 
from same fascicle. F-I, Same, from second specimen. J, Thoracic uncini. K, Uncini from first abdominal torus. L, Uncini from third 
abdominal torus. M, Anterior abdominal uncini from another specimen. N, Uncini from mid-abdominal torus (transitional region). O. 
Posterior abdominal uncini. P-T, Bayonet chaetae of one fascicle. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



57 




58 



T.G. PILLAI AND HA. TEN HOVE 



Table 5 S. ypsilon sp. nov. Measurements (mm) and counts. 



Total length 


27.5 


15.6 


14.1 


11.7 


6.3 


6.0 


4.6 


Thoracic width 


- 


0.6 


0.6 


0.4 


0.5 


0.5 


0.4 


No. of radioles 


- 


7/7 


8/8 


5/5 


6/6 


6/6 


4/4 


Abdominal length 


25.5 


11.0 


11.3 


8.5 


4.6 


4.3 


2.6 


No. of abdom. segs. 


113 


88 


131 


107 


37 


57 


37 


Capillaries from 


103 


80 


124 


99 


24 


44 


30 



Table 6 S. ypsilon sp. nov. Numbers of radioles. 



Table 9 S. ypsilon sp. nov. from Florida (EJ- 66-460). 

Measurements and meristic data from two longest specimens out 
of 22 measured. 



Total Thoracic 
length width 
(mm) (mm) 



Specimen 1 
Specimen 2 



No. of Abdomen 

radioles Length No. of Caps 
(mm) segs. from 



22.7 
20.7 



0.6 
0.8 



7/7 
6/6 



18.5 
16.7 



110 
130 



97 
111 



No. of specimens (n=37) 
No. of radioles (L/R) 



6 15 13 2 

7/7 6/6 5/5 4/4 



Table 10 Meristic and other data on S. ypsilon sp. nov. from 
Florida (EJ- 66-460). 



Table 7 S. ypsilon sp. nov. Numbers of thoracic chaetal tufts in 69 
specimens. 



No. ofspecim. 1 2 1 9 2 2 16 23 3 8 1 1 
Nos. of. tufts. 10/9 10/8 9/9 9/8 9/7 9/6 8/8 8/7 8/6 7/7 7/6 7/5 



Table 8 S. ypsilon sp. nov. Extent of thoracic membranes in 43 
specimens. 



Number of specimens 
Thor. membranes end on 



5 
6/6 



31 
6/5 



6 

5/4 



1 
5/3 



of the radioles (Fig. 6, E). Live material from Stns. 7.D03 
and 7.D05 showed a pair of pigmented ocelli at the base of 
each radiole, externally. An operculum is absent; there is a 
filamentous rudimentary operculumon each side (Fig. 6, E, 
L, O). 

Two clusters of reddish to reddish-brown prostomial ocelli 
are present. Although the width of the thorax ranges from 
0.4-0.6 mm in specimens preserved within their tubes, it can 
be wider in anterior portions of worms accidentally preserved 
outside their tubes (Fig. 6, F-H, L-O). The median lobe of 
the collar is sub-rectangular, rounded laterally and with a 
smooth mid-ventral notch, giving the entire collar a four- 
lobed appearance (Fig. 6, F-H & L-O). 

The numbers of thoracic chaetal tufts on the two sides 
range from 5 to 10, and may be symmetrical or asymmetrical, 
as are the endings of the thoracic membranes (Tables 7 & 8). 
Paired thoracic glands are absent. 

Collar fascicles bear up to about four fully developed 
bayonet chaetae and two more being formed deep within the 
fascicle, with a similar number of simple bladed chaetae. 
Juveniles possess fewer, often two fully developed bayonets 
and two more being formed within the fascicle. Each bayonet 
chaeta has a long serrated blade, a short unserrated notch, 
and few to several moderately large somewhat conical teeth 
with smooth tips on its basal boss (Fig. 7, A-E, F-I). The 
serrations of the blade are short towards its proximal part. 



No. of specimens 

(n = 14) 

No. of radioles 


4 
7/7 


2 
7/6 


4 
6/6 


1 
6/5 


2 
5/5 


1 

3/5 








No. ofspecim. 

(n = 29) 

No. of th. chaetal 


1 
10/9 


1 

10/7 


1 
9/9 


4 
9/8 


6 

8/8 


9 

8/7 


4 

7/7 


2 
7/6 


1 

7/5 


tufts 




















No. of specimens 

(n = 20) 

Thor. membrane 


1 

7/5 


1 

6/5 


10 

5/5 


7 
5/4 


1 

4/4 










ends 





















but somewhat pilose distally. Thoracic uncini are mostly with 
6 teeth, but some have 4 or 5 (Fig. 7, J). Anteriorly there are 
up to 4 flat trumpet chaetae in each bundle, posteriorly there 
are 1 or two capillaries instead. Anterior abdominal uncini 
usually have 4 or 5 teeth arranged in a single row (Fig. 7, 
K-M). The posterior abdominal uncini are rasp-shaped, with 
a single anterior tooth and several (4-6) rows of teeth 
posterior to it (Fig. 7, O). In between, there is a progressive 
reduction in the number of teeth in a single row, and a 
corresponding increase in the rasp-shaped area (Fig. 7, N). 
The special adaptations of the body of the worm in relation to 
the internal structures of the tube are as follows: A narrow 
longitudinal groove extends along the mid-dorsal line of the 
abdomen and thorax (Fig. 6, C, D, H-K, O). The abdomen 
and thorax are also grooved ventrally, and within this longitu- 
dinal groove, forms a cord-shaped longitudinal ridge (Fig. 6, 
C, D, I-L). The orientation of the worms within their highly 
coiled tubes is such that the dorsal groove is applied to the 
serrated dorsal ridge of the tube, and the cord-shaped ventral 
abdominal ridge fits into the gutter-shaped part of the 
Y-shaped ventral ridge of the tube. The latter, in turn, fits 
into the ventral groove of the body. 

Collections from the western Atlantic. 5. ypsilon has 
also been collected from Florida, Bermuda and Aruba. 

Study of material from Florida (EJ- 66-460) provided the 
following data: The external diameter of the tubes attains 
1.1 mm. A granular overlay is present. The external coloura- 
tion varies from faintly creamish to faintly pinkish. Their 
internal colouration and structures are similar to those from 



Fig. 8 Spiraserpula paraypsilon sp. nov. A, Tube from Curasao, NA, Cornelisbaai, showing granular overlay and longitudinal ridges. B-N. 
From Klein Bonaire Stn. 2105A. B, Tube with indistinct longitudinal ridges. C-N, From holotype. C, Tube fragment showing serrated 
dorsal ridge along convex wall. D, Tube fragment with Y-shaped ventral ridge along opposite wall. E & F, Adult worm; E, Radioles of 
both sides, with very long pinnule-free tips and lacking rudimentary opercula. F, Body showing dorsal and ventral longitudinal abdominal 
grooves and ventral pigment patches. G-L, Bayonet collar chaetae. M, Thoracic uncini of holotype showing lateral denticles. N, Anterior 
abdominal uncini, with more prominent denticles. 






ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



59 




60 



T.G. PILLAI AND HA. TEN HOVE 



the Cape Verde Islands. The colouration of the worms in 
alcohol shows a difference. The radioles and body have an 
overall fleshy to dark reddish-brown colour, with blackish 
pigment clusters ventro-laterally in the abdominal segments. 
Whether this colour difference is due to different methods of 
fixation or not needs to be verified. Maximum sizes encoun- 
tered have been given in Table 9. 

An operculum is absent, but a pair of rudimentary opercula 
is present. Up to 10 thoracic chaetal tufts per side were 
counted, and the extent of the thoracic membranes is vari- 
able, however, never reaching the last thoracic chaetiger. 
Meristic and other data are given in Table 10. 

The collar chaetae (Fig. 6, P-T), are larger than those of 
the specimens from the type locality, but otherwise similar. In 
the abdomen, up to 10 flat trumpet shaped chaetae per 
fascicle were counted. 

The other two Florida samples are similar. However, in 
sample EJ 67-76, one specimen lacking radioles has 12 
thoracic chaetigers on the left and 11 on the right, with the 
thoracic membranes ending on the 6th and 5th chaetiger, 
respectively. The samples from Bermuda and Aruba are 
similar to those from Florida. 

Live material. There are some intra-specific colour varia- 
tions inS. ypsilon sp. nov., as observed in collections from 
different stations in the Cape Verde Islands: 

Stn.6.D02: Branchial radioles distally orange, proximally 
pinkish orange, except for a bright red spot where the 
radioles meet. Thorax is transparently reddish, with two 
subcutaneous brown spots laterally. Abdomen is pink or 
orange, with brown sides. 

Stn.7.D03: Distal half of the short radioles banded white 
and yellowish orange, basal half bright red. Basal radiole 
parts with oval lens-shaped structures, apparently ocelli. 
However, after preservation no lenses could be found in this 
material, not even after staining in methylene blue. Thorax 
bright red, abdomen orange, with brownish-green granules 
laterally. 

Stn.7.D05: Radioles transparent, hyaline, with a single row 
of pigmented spots at the base. No lenses visible, even with a 
compound microscope. Thorax and abdomen orange-brown. 
Sides of abdomen show brown granules (in four specimens). 
In two other specimens the radioles are hyaline, with trans- 
verse orange bands and red pigment spots at their bases. The 
rest of the body is red, otherwise similar to the other four 
specimens. 

Etymology. The specific name refers to the Y-shaped 
internal ridge. 

Habitat and distribution. As revealed by several scuba 
dives and littoral surveys in the Cape Verde Islands, 5. 
ypsilon sp. nov. occurs on various hard substrata in depths 
occupied by S. massiliensis (Zibrowius) in the Mediterra- 
nean. At Stn. 6.D02, for instance, the ceiling of a half metre 
deep cave, at a diving depth of 14m, was covered with crusts 
of partially erect tubes of 5. ypsilon. However, 5. massiliensis 



did not turn up in any of the collections from the Cape Verde 
Islands. The single, eroded tube of S. ypsilon sp. nov. from 
the dredging station 6.148 (100-200 m) was probably trans- 
ported down the slope. 

Although the Western Atlantic and Cape Verde Islands 
material show some differences, they appear inadequate to 
separate them into distinct species. The Western Atlantic 
range is from Bermuda to Aruba. A species from the 
Caribbean which has similar internal tube structures, but 
differs in other respects, is described next. 

Spiraserpula paraypsilon sp. nov. 
(Figs.8, A-N; 9, A-R; 3, G) 

Material examined. 

Bonaire (Neth. Ant.): 1. Klein Bonaire, N, half mile E of 
Westpunt, reef, little sand, corals, 38 m, legit H.A. ten Hove, 
l.vii.1970, Stn. 2105A, HOLOTYPE & PARATYPE 2. 
ZMA V. Pol.3714; PARATYPE 1, BM(NH) 1992.156. 2. 
Lac, dam, pool in wash of plunging breakers, Diploria, 
Millepora, Pontes, cobble in coarse sand, 50 cm, from corals, 
legit H. A. ten Hove, 15.vii.1970, Stn. 2122C, 
(PARATYPES 6-8, ZMA V.Pol. 3717). 3. Plaja Frans, on 
dead coral covered with calcareous algae, little sand, 
1.0-1.5 m, legit H. A. ten Hove, 16.vii.1970, Stn. 2110A 
(portions of tube, 1 incomplete worm, BM(NH) 1992.157). 4. 
Karpata, steep reef, drop off and flat above, 15^ m, from 
living corals, legit H.A. ten Hove, 19. v. 1987, Stn. 87-5 
(PARATYPES 4-5, USNM 130992). 

Curacao (Neth. Ant.): 5. Cornelisbaai, sandy reef, from the 
underside of dead plate-shaped coral, 15 m, legit H. A. ten 
Hove, 15.xi.1988 (PARATYPE 3 USNM 130991) and 17.i. 
1990 (4 specimens, AM W20338). 6. Piscaderabaai, outer bay 
in front of Carmabi, rubbish on reef, 10 m, legit H. A. ten 
Hove, 10-12.U990 (19 specimens, ZMA V.Pol.3718, 
BM(NH) 1992.158-165, FSBC I 39196. 7. Salinja Fuik, reef 
in front, 20-25 m, legit H. A. ten Hove, 18.U990 (25 
specimens, NSMT). 

Type locality. Klein Bonaire, Curacao. 

Description. 

tubes: White, flattened, and with a granular overlay. The 
maximum external diameter of the tube of the holotype is 
2.0 mm. A median and about 3 pairs of lateral longitudinal 
ridges can be observed (Fig. 8, A). In an empty tube from the 
type locality with a diameter of 1.5 mm, the number of ridges 
is less distinct (Fig. 8, B). ITS, located within the first formed 
coiled parts, are translucent white, and very similar to those 
5. ypsilon. They consist of a serrated dorsal ridge along the 
convex wall (Fig. 8, C) and a Y-shaped ventral ridge along the 
opposite side (Figs.8, D; 3, G). The serrations of the dorsal 
ridge are pointed and directed somewhat posteriorly. Tubes 
found on asbestos plates (Piscaderabaai) were clearly branch- 
ing, as described in detail for S. caribensis (Fig. 16, A & B). 
worms: The total length of the holotype is 16.4 mm. With a 
thoracic width of 0.8mm, it is stouter than S. ypsilon. The 



Fig. 9 Spiraserpula paraypsilon sp. nov. A-N, From Klein Bonaire Stn. 2105A; O-R, from Bonaire, Karpata Stn. 87. 5. A-J, From juvenile 
paratype. A, Tube showing start of external longitudinal ridge and shallow transverse growth markings. B, Posterior tube fragment with 
serrated dorsal ridge. C, Radioles with long pinnule-free tips, a rudimentary operculum on the left and none on the right. D & E, Two 
views of body showing pigment patches in both, dorsal an ventral longitudinal abdominal grooves (D), and extent of thoracic membrane 
(E). F-J, Bayonet chaetae, including one newly formed within the fascicle (J). K-N, From holotype. K & L, Tube fragments (K) and the 
other from a more posterior coil, with serrated dorsal ridge. M & N, Two views of body showing pigment patches, dorsal and ventral 
longitudinal abdominal grooves (M), and extent of thoracic membrane (N). O-R, Bayonet chaetae of one specimen. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



61 




62 



T.G. PILLAI AND H.A. TEN HOVE 



abdomen is 9.9mm long, with about 98 segments, and capil- 
laries in the last 20. 

The pinnule-free tips (Fig. 8, E) are quite conspicuous and 
much longer than those of S. ypsilon, being up to about 
1.9 mm. They constitute nearly half to more than half the 
length of the radioles (about 3.7 mm). The number of 
radioles, 11 on each side, is higher than that of S. ypsilon 
(maximum 8). They bear at intervals what appear to be 
narrow, transverse, lenticular lacunae. Two pigmented pros- 
tomial ocellar clusters are present. 

The rest of the body (Fig. 8, F) is similar to that of S. ypsilon 
in many respects. In alcohol, the abdomen has an overall 
pinkish colour, with clusters of reddish-brown pigmented 
cells ventro-laterally. 

Thoracic chaetigers number 7 on both sides. Thoracic 
membranes end on the third thoracic chaetiger (second 
uncinigerous segment) on both sides of the thorax. Paired 
ventral thoracic glands were not seen. 

A collar fascicle of the holotype has five fully formed 
bayonet chaetae, and a developing one deep within. A 
paratype from Karpata has 6 fully formed bayonets and one 
newly forming one within the fascicle. Each bayonet chaeta 
consists of a long, narrow serrated blade, and a considerably 
expanded basal boss bearing several moderately large, some- 
what pointed teeth (Fig. 8, G-L). The serrations are short and 
fine proximally, but longer and pilose distally. The unser- 
rated notch may be very short, up to about twice the length of 
the longest teeth, or almost lacking (Fig. 8, I). The thoracic 
and anterior abdominal uncini bear 5 teeth in a single row. 
They differ from all the other known species of the subgenus 
in possessing minute denticles on their sides (Fig. 8, M,N). 

The adaptations of the worm in relation to ITS are similar 
to those of S. ypsilon. The dorsal longitudinal abdominal 
groove is applied to the serrated dorsal ridge of the tube, and 
the Y-shaped ventral ridge is enclosed within a ventral 
abdominal groove. Within the latter, a cord-shaped abdomi- 
nal ridge fits into the gutter-shaped part of the Y. 

The paratype from the type locality is a juvenile. Its tube 
(Fig. 9, A) shows faint transverse grooves mainly, but the 
beginnings of a granular overlay and longitudinal ridges can 
also be seen. ITS and adaptations of the body are identical to 
those of the holotype (Fig. 9, B,K,L). Measurements and 
counts of the worm are as follows: Length 6.2 mm, thoracic 
width 0.5 mm, radiolar length 2.1 mm, pinnules 1.1 mm, 
abdomen 4.2 mm, 53 segments, with capillaries on the last 
10. The number of thoracic chaetigers, ending of the thoracic 
membranes (Fig.9,E), and colour, are the same as in the 
holotype. 

The 7 pairs of radioles already approach the maximum 
number in other material, except the holotype, and their long 
pinnule-free tips are similar to those of the holotype (Fig. 9, 
C). However, there is a very short and slender rudimentary 
operculum on one side, while it is lacking on the other (Fig. 9, 
C), indicating that both may become completely lost in older 
specimens (holotype). A similar condition is found in one of 
the specimens from Karpata (below). 

Collar fascicle with four fully formed bayonet chaetae 
(Fig. 9, F-I) and a developing one deep within (Fig. 9, J). 
Their basal bosses are not as expanded as in the holotype and 
the blades are shorter. The uncini are similar to those of the 
holotype. 

The tubes of the three specimens from Karpata agree with 
those from the type locality in being white externally, pinkish 
internally, and bearing the Y-shaped ventral ridge and ser- 



rated dorsal ridge. The serrations of the latter bear posteri- 
orly directed tapered tips (Fig. 9, K,L). The radioles of all 
specimens are detached, highly contracted, and do not clearly 
show the extent of the pinnule-free tips. One crown has a 
short rudimentary operculum on each side, the second has a 
rudimentary operculum on one side but lacks it on the other, 
and the third half crown has a rudimentary operculum which 
is very reduced and filamentous. The thoracic width ranges 
between 0.5 mm and 0.7 mm. The abdomen of the longest 
specimen is 10.8 mm long and has about 59 segments, with 
capillaries on the last six; that of the shortest is 7.3 mm, but 
has about 86 segments, with capillaries on the last six. The 
numbers of radioles, thoracic chaetal tufts and the extent of 
the thoracic membranes in the three specimens is provided in 
Table 11. 

In two specimens the broad thoracic membranes are folded 
outwards against the sides of the thorax (Fig. 9, M,N). 
Bayonet collar chaetae (Fig. 9, O-R) are similar to those of 
the holotype, but lack an unserrated notch. Thoracic and 
anterior abdominal uncini are also similar to those of the 
holotype, with 4 or 5 teeth in a single row. Flat trumpet 
chaetae number 6-8 in a bundle. Preserved in alcohol, the 
abdominal segments show clumps of reddish-brown pig- 
mented cells ventrolaterally, and of larger yellowish or orang- 
ish cells ventrally (Fig. 9, N). 

The tube of the single juvenile paratype from Curacao 
resembles that of the holotype; its diameter is 1.5 mm. The 
worm has 6 radioles on the left and 5 on the right. As in the 
specimens from Bonaire, the pinnule-free tips are very long. 
However, both rudimentary opercula have already been lost 
at this stage. The thorax is 0.5 mm wide and has 7 pairs of 
chaetigers. The thoracic membranes end on the third thoracic 
chaetiger. Two clusters of prostomial ocelli are present. 
Thoracic glands were not seen. The chaetae also agree with 
those of the specimens from Bonaire. In recently collected 
material (Curacao, Piscaderabaai, 10. i. 1990), thoracic mem- 
branes end at 4/5, 4/4 respectively; the pinnule-free tips 
generally are very long, rarely short; rudimentary opercula 
are present in four specimens, absent in three. 

Live material. According to the field notes, rudimentary 
opercula could not always be found, even in living specimens 
from Curasao with radioles extended. The colouration of the 
radioles is somewhat variable, often (transparently) whitish 
to creamish, rarely yellowish to slightly orange or even 
completely hyaline. At the base of the radioles there is a 
series of up to six pairs of reddish spots, absent however in 
the dorsal- and ventralmost radioles. Body predominantly 
orange-brownish with up to 15 greenish-brown granules per 
segment ventro-laterally in the abdomen and dorsally in the 
thorax. 

Etymology. The specific name paraypsilon indicates the 
close resemblance of the species to S. ypsilon. 

Habitat and distribution. Occurs in shallow, clear, oce- 

Table 11 S. paraypsilon sp. nov. Meristic and other data from 
specimens. 



Specimen nos. 

No. of radioles 

No. of thoracic chaetal tufts 

Thoracic membrane ends 



1 


2 


3 


8/? 


8/8 


8/7 


?/? 


8/9 


7/7 


?/? 


3/3 


3/3 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



63 




U V 



Fig. 10 Spiraserpula singularis sp. nov. From type series. A & B, Two tubes, the second one younger. C, Substratal view of first formed coil 
showing V-shaped dorsal ridge; a ventral ridge is absent. D, Same, smaller specimen. E, Paratype. F & G, Holotype and radioles. H-L, 
From whole mount of another paratype: H & I, Two views of worm; J, only available bayonet chaeta; K, L, thoracic uncini; M, anterior 
abdominal uncini. M-R, From whole mount of another specimen: N, anterior abdominal uncini and flat trumpet chaetae; O-R, Bayonet 
collar chaetae, two present per side. S-V, Bayonet chaetae from whole mount of third specimen. 



64 



T.G. PILLAI AND HA. TEN HOVE 



anic waters, at depths of up to 40 m on coral reefs. Hitherto 
collected only from Bonaire, and Curasao, in the Caribbean. 

Spiraserpula singularis sp. no v. 

(Figs. 10, A-V; 3, B; P1.2, E & F) 

Material examined. 

Puerto Rico : 1. Isla Matei, near buoy of Marine Institute, 
vertical reef with surge channels, no sand, from living corals, 
29-33 m, legit H. A. ten Hove, 2.x. 1970, Stn. 2136A, 
(HOLOTYPE & 3 PARATYPES, ZMAV.Pol.3710). 
Curacao, (Neth. Ant.): 2. Salinja Fuik, reef in front, marine 
park, 20-25 m, legit H. A. ten Hove, 18. i. 1990 (2 specimens 
BM(NH) 1992.166-167). 3. Piscaderabaai, outer bay, W of 
entrance, sandy reef, 20 m, from underside of coral debris, 
not in sediment, legit H. A. ten Hove, 12. i. 1990 (3 speci- 
mens, BM(NH) 1992.168-170). 

Type locality. Puerto Rico. 

Description. 

tubes: White, very tiny, one of the smallest species in the 
genus. They may occur in mutually bonded aggregations of a 
few individuals, or singly. Their coil diameters range from 
1.2-1.3 mm. A granular overlay is present (Fig. 10, A,B), 
which makes the external diameters of the tubes 
(0.5-0.6 mm) considerably larger than their internal diam- 
eters (0.2-0.25 mm). Their apertures bear small, somewhat 
lobed, peristome-shaped extensions (Fig. 10, A,B), similar to 
those found in S. massiliensis. 

ITS consist of a V-shaped dorsal ridge, actually an inverted 
V, along the convex side of the first formed coil (Figs. 10, 
C,D; 3, B). The two arms of the V are broader and outwardly 
curved posteriorly, and their edges are smooth. Anteriorly 
the dorsal ridge is a smooth plate only. A ventral ridge is 
absent. When the worm is withdrawn into the tube, the 
posterior, mid-dorsal part of the abdomen is applied to the 
dorsal ridge. 

worms: Four specimens were taken out of their tubes 
(Fig. 10, E-I). The holotype (Fig. 10, F), which is the largest, 
is only 5.7 mm long, 0.2 mm wide in the thorax, and its 
abdomen is 4.6 mm long. There are four pairs of radioles 
which, including the short and slender pinnule-free tips 
(Fig. 10, G), are about 0.55 mm long. There is a rudimentary 
operculum on each side. Radioles are missing in the other 
three specimens. However, a detached operculum was found 
in the vial containing the specimens, and it is not certain 
whether it belongs to one of them or another species. 

Two clusters of prostomial ocelli are present. Five or six 
globular ventral thoracic glands are present, more or less 
arranged in a V. The numbers of thoracic chaetigers on the 
two sides in the four specimens are: 9/9, 9/8, 8/8, and 7/7. It 
was not possible to establish the extent of the thoracic 
membranes due to the extremely small size of the worms. An 
apron is, however, absent. One paratype with an abdominal 
length of 1.95 mm has 29 segments, with capillaries on the 
last 5, and another 3.0 mm long with 39 segments, but the 
capillaries cannot be seen, having probably been damaged. 

There are two bayonet chaetae in each collar fascicle. They 
have moderately long serrated blades and 2-A teeth on the 
basal boss and some accessory ones (Fig. 10, J, O-V; PI. 2, E). 
The unserrated notch is 1/5 the length of the blade. Thoracic 
uncini (Fig. 10, K,L) and anterior abdominal uncini (Fig. 10, 
M,N) have 6 and 4-6 teeth, respectively, all in a single row. 
The middle abdominal uncini are rasp-shaped (PI. 2, F), with 



up to 3 transverse rows of teeth above the single anterior 
tooth. The abdominal segments bear 1 or 2 flat trumpet 
chaetae in each bundle; one side is thickened into a claw- 
shaped process (Fig. 10, N). 

Remarks. In the comparison with other Caribbean species, 
S. singularis would key out mainly on the absence of a ventral 
longitudinal ridge/row of teeth and probably also the absence 
of an operculum. So far, the presence of an operculum has 
been observed only in a doubtful field identification. The 
form of thoracic glands, shape of dorsal ridge and collar 
chaetae are similar to those in S. plaiae. 

Etymology, singularis (Latin) = unique; referring to the 
unique ITS. 

Habitat and distribution. S. singularis sp. nov. appears 
to be a shallow water coral reef dweller. It has hitherto been 
collected only from Puerto Rico and Curacao. 

Spiraserpula karpatensis sp. nov. 

(Figs.ll, A-K;3,N) 

Material examined. 

Bonaire (Neth. Ant.): 1. Karpata, reef, 10 m, cryptic, legit H. 

A. ten Hove, 9.xi.l988 (HOLOTYPE, ZMA V.Pol.3712; 

PARATYPE, BM(NH) 1992.171). 

Curacao (Neth. Ant.): 2. Reef in front of Salinja Fuik, buoy 

13 of Marine Part, 20-30 m, corals and sandy/silty areas in 

equal amounts. From under side of coral debris, not in 

sediment, legit H. A. ten Hove, 18. i. 1990 (1 specimen, ZMA 

V. Pol. 3875). 

TYPE LOCALITY. Bonaire (Netherlands Antilles). 

Description. 

tubes: Pink, quite small, and lack longitudinal ridges. A pink 
translucent granular overlay is present (Fig. 11, A). The tubes 
of both the types are coiled upon themselves, one much more 
than the others (Fig. 11, D). One has an erect part 2.0 mm 
long, and a funnel-shaped, outwardly curved peristome 
(Fig. 11, A), while the other has a somewhat thickened 
anterior end (Fig. 11, D,E). The pink colouration gradually 
fades to white towards the anterior end. The diameter of the 
tubes is 0.6-0.7 mm in the attached parts, 0.4-0.6 mm in the 
erect parts. 

ITS consist of a serrated ventral ridge and an unserrated 
dorsal ridge, with a sharp edge in cross-section (Fig. 3, N). 
The dorsal ridge may be absent (Fig. 11, B,C), or greatly 
reduced (Fig. 11, D). In the latter it can be seen as a short 
crescentic ridge through the broken end of one of the coils. 
The serrated ventral ridge is regularly present (Fig. 11, B, C, 
and bottom left of D). 

worms: The holotype is incomplete posteriorly (Fig. 11, F), 
broken in three parts, with a total length of 4.3 mm. An 
operculum and 4 radioles are present on the left side; the 
radioles on the right are missing. The length of the radioles is 
approximately 0.8 mm, with a pinnule-free tip of 0.1 mm. 
The operculum (Fig. 11, G,H), is 0.26 mm long, and 0.28 mm 
in diameter; inclusive of peduncle it is about 1 mm long. 
Although bell-shaped, it is slightly zygomorphic, and has 
numerous fine lobes, similar to that of S. plaiae described in 
this paper. Branchial eyes have not been observed in the 
fresh material. Two clusters of prostomial ocelli are present. 
The thorax has 8 chaetigers on each side. The thoracic 
membranes extend to the third chaetiger on the left (Fig. 11, 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



65 




Fig. 11 Serpulu karpalensis sp. nov. A, Tube with granular overlay, erect part and funnel-shaped peristome. B & C, Substrata! view of two 
tubes, opened to show internal serrated ventral ridge along convex wall, but absence of dorsal ridge. D, Aggregation of tubes, some 
sections showing a very short crescentic dorsal ridge. E, Erect part from same aggregation showing somewhat thickened distal end; granular 
overlay. F, Anterior end of holotype showing collar and thoracic membrane. G & H, Two views of zygomorph operculum. I-K, Bayonet 
chaetae. 



F) and the fourth on the right. It is not certain whether 
ventral thoracic glands are present, but see note on live 
material below. 

Each collar fascicle bears 3 bayonet chaetae (Fig. 11, I-K), 
with moderately long, finely serrated blades, a moderately 
long unserrated notch, and 3 teeth on the basal boss; the third 
tooth may sometimes be difficult to observe and may be 
reduced to a scar. Thoracic uncini have 6 (exceptionally 7) 
teeth, anterior abdominal uncini 5, arranged in a single row. 
The middle abdominal uncini are rasp-shaped, with 3-5 teeth 
above the single anterior tooth. At least 35 abdominal 
chaetigers are present, the last 7 with capillary chaetae. 
Abdominal flat trumpet chaetae number 2-3 per bundle. 

The specimen from Curasao agrees in most details with the 



type material. Its numbers of radioles are 5/5, a long filamen- 
tous rudimentary operculum is present opposite the opercu- 
lum, and it has 38 abdominal chaetigers. 

Live material. As observed in material collected in 1990, 
radioles are transparently lemon. Thorax ventrally with 5 
bright red globules arranged in a V, presumably thoracic 
glands. 

Etymology, named after the type locality, the coral reef in 
front of the Sentro Ekologiko, Karpata. 

Habitat and distribution. A shallow water cryptic reef 
dweller. Has hitherto been recorded only from its type 
locality in Bonaire, and Curacao. 



66 



T.G. PILLAI AND HA. TEN HOVE 




Fig. 12 Spiraserpula zibrowii sp. nov. From type specimens. A & B, Substratal view of two tubes opened to show the unserrated dorsal 
ridge. The serrated ventral ridge consists here of a row of isolated teeth, but is a continuous ridge in the remaining material. C, Juvenile 
paratype. D, Older specimen (holotype) broken in two. Anterior part with right rudimentary operculum; posterior abdomen with part of 
dorsal ridge attached to mid-dorsal groove. E, Paratype. F & G, Two bayonet chaetae from holotype; H & I, two bayonet chaetae from 
paratype. J-O from paratype. J, Thoracic uncini of small specimen with single row of teeth anteriorly and a cluster of more than one row 
posteriorly. K, Similar uncini from first abdominal torus. L, Uncini from second abdominal torus; there are less teeth in a single row. M & 
N, Anterior abdominal uncini, and flat trumpet chaetae with large lateral tooth. O, Posterior abdominal uncini; except for a single anterior 
tooth, the uncini are rasp-shaped, with teeth in several rows. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



67 



Spiraserpula zibrowii sp. nov. 
(Figs. 12, A-N; 3, O; P1.4, A-D) 

Material examined. 

Curacao (Neth. Ant.): 1. Lagoon of San Juan, E, raised reef, 
lagoon side, Halimeda present, limestone cobbles, 10-15 cm; 
from up to 20 cm deep crevices between cobbles, legit H. A. 
ten Hove, 29.vi.1970, Stn. 2043 [HOLOTYPE & 
PARATYPES 3-5: ZMA V.Pol. 3707; PARATYPES 1 
(slide), 2 (worm & tube fragments), and 7 (empty tube: 
BM(NH) 1992.148-150); PARATYPE 6: USNM 130980 
(unopened tube)]. 

Bonaire (Neth. Ant.): 2. Kralendijk, Flamingo Beach Hotel, 
from corals, partly in sand, 45 m, legit H. A. ten Hove, 
27.vii.1970, Stn. 2115D, (4 empty tubes, BM(NH) 
1992.151-155). 3. 250 m N of Witte Pan, sandflat below reef, 
47 m, mainly from the side of boulders, partly buried in sand, 
legit H. A. ten Hove, 3.vii.l970, Stn. 2117B (4 empty tubes, 
ZMAV. Pol. 3708). 

Type locality. Curasao (Netherlands Antilles). 

Description. 

TUBES: Whitish, very tiny, and coiled upon themselves like 
spirorbids, either individually (Fig. 12, B), or in mutually 
bonded aggregations of a few individuals. The direction of 
coiling may reverse (see below). A fine granular overlay is 
present. Longitudinal ridges are absent, but fine, smooth, 
transverse growth markings are present. Juvenile tubes are 
white. Although older tubes are white posteriorly, they have 
a greyish-brown overlay anteriorly. The diameter of an 
individual coil is 0.73 mm, with a tube diameter of 0.18 mm. 
The maximum tube diameter is only 0.44 mm, which is the 
smallest among the known species of the genus. 

ITS consist of a serrated ventral ridge and an unserrated 
dorsal ridge (Fig. 12, A,B). The ventral ridge may consist 
either of a continuous row of serrations, or only of a short 
row of small separate teeth (Figs. 12, A,B; 3, O). The dorsal 
ridge is colourless and transparent, wedge- to Y-shaped in 
cross-section, with its edges curved in places; it is spiral on a 
columella-shaped axis when the tube is coiled upon itself. 
Lateral ridges have not been found. The interior of the tube 
may have a creamish lining. 

One tube is coiled in one direction proximally, and in the 
opposite direction distally. In the proximal coil the ITS are 
similar to those described above. However, the distal coil has 
only a columella-shaped axis with a dorsal ridge, which 
became detached from the tube and is shown in situ (Fig. 12, 
E); a serrated ventral ridge is absent here. 

The mid-dorsal and mid-ventral longitudinal grooves of the 
abdomen are applied to the unserrated dorsal ridge (Fig. 12, 
D) and serrated ventral ridge of the tube, respectively. 
WORMS: Measurements and meristic data are presented in 
Table 12. 

The right branchial half of the holotype (left missing) 
shows a rudimentary operculum. The latter is present on both 
sides in the first paratype, but not in the second which is a 
juvenile. The numbers of radioles on both sides are ?/3, 4/4 
and 3/4, respectively. The pinnule-free tips are about 1/5-1/7 
of their total length. 

Two clusters of prostomial ocelli are present. The numbers 
of thoracic chaetal tufts on both sides in the three specimens 
are: 7/7, 8/7 and 8/7, respectively. The thoracic membranes 
end on chaetigers 3/3 in the holotype and 4/4 in the juvenile 
paratype; they are damaged in the second paratype. Two 



Table 12 S. zibrowii sp. nov. Measurements and meristic data 
from Holotype and two paratypes. 





Total 
length 
(mm) 


Thoracic 
width 
(mm) 




Abdomen 






Length 
(mm) 


Number of 
segments 


capillaries 
on 


Holotype 
Paratype 1 
Paratype 2 


9.7 
7.0 
3.4 


0.23 
0.18 
0.18 


8.5 
5.8 
2.1 


54 
43 
27 


4 
7 
9 



translucent ventral thoracic glands are present, although not 
as easily discerned as in some of the other species. 

There are 2 or 3 fully developed bayonet chaetae per side 
in the juvenile, 4 in the older specimens. They have moder- 
ately long serrated blades, an unserrated notch which is about 
1/4-1/5 the length of the blade, and 4 or 5 somewhat large 
teeth and some accessory ones on the basal boss (Fig. 12, F,I; 
PI. 4, A). The thoracic uncini have a single row of 6-7 teeth 
(PI. 4, B). Anterior abdominal uncini bear a cluster of small 
teeth in two to seven rows at their posterior ends, and a single 
row of larger teeth anteriorly; this type of uncini may occur in 
juvenile specimens also (Fig. 12, J-M). The posterior abdomi- 
nal uncini are, however, similar to those of the other species 
in being rasp-shaped, with 6 transverse rows of 2-5 teeth 
each, except for the single anterior tooth (Fig. 12, O; PI. 4, C). 
The abdominal flat trumpet chaetae number about 5 per 
bundle. Their somewhat triangular, curved distal ends are 
thickened and hooked at one end, and drawn out into an 
acute angle at the other (Fig. 12, N; PI. 4, D). Up to 54 
abdominal segments are present, the last 4-9 with capillary 
chaetae. 

Remarks. The collections from Bonaire, a mere 50 km from 
Curasao, consist of a total of 8 empty tubes whose ITS are 
identical with those of the present species. The largest tube 
from Witte Pan has a coil diameter of 2.2 mm; two tubes have 
erect portions with peristomes. Three tubes are white. The 
fourth is creamish in colour, with a creamish interior lining. 
Fine transverse growth markings are present on all. The 
serrations of the ventral ridge are arranged on a low longitu- 
dinal ridge in some of them. 

In the absence of worms, and the markedly different 
habitat from which they were collected (at a depth of 
45-47 m), these tubes cannot be conclusively identified as S. 
zibrowii. 

Etymology, named after H. Zibrowius, who recognized 
some of these small species as being new. 

Habitat and distribution. Appears to be a shallow water 
species inhabiting crevices between boulders and their under- 
sides in sandy areas close to coral reefs. Hitherto collected 
from Curasao. Two uncertain records from Bonaire. 



Spiraserpula plaiae sp. 
(Figs. 13, A-T; 3, K) 



nov. 



Material examined. 

Curasao (Neth. Ant.): 1. Salinja Fuik, near Ceru Preekstul, 
open reef, coral debris, 33 m, from limestone boulder on 
sand, legit H. A. ten Hove, 18. ix. 1970, Stn. 2088A (HOLO- 
TYPE & PARATYPES 1 & 5: ZMA V. Pol.3713; 



68 



T.G. PILLAI AND HA. TEN HOVE 



PARATYPES 2, 4, & 6: BM(NH) 1992.173-174; 
PARATYPE 3: USNM 130990). 2. Reef in front of Salinja 
Fuik, buoy 13 of marine park, coral debris, 18-27 m, legit H. 
A. ten Hove, 18. i. 1990 (4 specimens, ZMA V. Pol. 3874). 3. 
Cornelisbaai, E, steep reef, coral debris, 18-26 m, legit H. 
A. ten Hove, 17. i. 1990 (6 specimens, ZMA V. Pol. 3873). 4. 
Piscaderabaai, outer bay W of entrance, sandy reef, coral 
debris, legit H. A. ten Hove, 12. i. 1990 (5 specimens, ZMA 
V. Pol.3872). 

Type locality. Curacao (Netherlands Antilles). 

Description. 

tubes: White to greyish-brown, occurring either individually 
coiled upon themselves (Fig. 13, A), or in mutually bonded 
aggregations of a few individuals (Fig. 13, B). They are 
sub-circular in cross-section, with faint lateral ridges, and 
bear fine smooth transverse ridges, and often have erect 
anterior ends (Fig. 13, A,B). A fine opaque granular overlay 
is present (Fig. 13, B), which can be seen under special 
illumination only. Their external diameter attains 1.0 mm, 
their erect portions somewhat smaller. The inside of one tube 
has a light caramel coloured lining. 

ITS consist of a serrated ventral ridge along the concave 
side of the tube (Fig. 13, C-E, H), which may not be well 
developed and represented only by a few isolated or coa- 
lesced teeth slanting backwards (Fig. 13, F), and a smooth 
dorsal ridge on the convex side (Figs. 13, F,G; 3,K). The 
latter is wedge-, T to Y-shaped in cross-section. In specimens 
coiled upon themselves, the dorsal ridge occurs spirally on a 
columella-shaped axis (Fig. 13, I). A short accessory latero- 
dorsal ridge, which tapers anteriorly and posteriorly, may 
also be present on either side (Fig. 13, F). Their edges are 
unthickened. 

In life, the mid-ventral and mid-dorsal longitudinal grooves 
of the abdomen are applied to the serrated ventral and 
smooth dorsal ridges, respectively, of the tube (Fig. 13, G). 
worms: Six specimens were taken out of their tubes. The 
abdomen is complete in only one. Even though preserved in 
alcohol, the abdominal segments still show clusters of pig- 
mented specks laterally, light yellowish in one specimen, light 
to bright orange in two, light to dark brown in two, and 
uniformly caramel coloured in another. 

Five specimens have an operculum on one side and a 
rudimentary operculum on the other; the branchial crown is 
partly missing in the sixth. The length of the operculum and 
peduncle varies from 1.0 mm in a juvenile paratype to 
1.8 mm in the holotype. The operculum itself is 0.4—0.5 mm 
long, 0.3-0.5 mm wide. It is zygomorph, attached to the 
peduncle eccentrically, and bears numerous (up to 50) radii 
(Fig. 13, K-M). The distal diameter of the peduncle is 1/3 to 
2/3 that of the opercular base. The numbers of radioles on 
both sides are 6/5, 5/4, 4/5, 4/4 and 4/3. They end in short 
slender pinnule-free tips, which are about 1/5 to 1/7 the length 
of the radioles (Fig. 13, K). The only complete specimen 
(Fig. 13, J), is 4.9 mm long, with 42 abdominal segments, the 
last 10 with capillaries. However, in another specimen, which 



is incomplete (Fig. 13, M), 76 abdominal segments could be 
counted. 

Two clusters of prostomial ocelli are present. The num- 
bers of thoracic chaetal tufts are 11/7, 9/9, 9/8, 8/8, 6.6. 
Thoracic membranes extend to chaetigers 6/5, 4/4, 4/3, 4/?, 
and they are damaged on both sides in the fifth. Two 
groups of transparent to translucent ventral thoracic 
glands, arranged in a V, and of unknown function, are 
present. Further studies are needed to find out if they 
could be responsible for secreting the caramel coloured 
inner lining of the tube. 

Collar fascicles bear 2 or 3 fully formed bayonet chaetae 
each, and a newly formed one deep within the bundle. Each 
bayonet chaeta consists of a moderately long blade, a moder- 
ately long unserrated notch which is 1/3 to 1/4 the length of 
the blade, and 2-4, seldom 5, teeth on the basal boss (Fig. 13, 
N-T). The teeth are comparatively larger as their number 
decreases (Fig. 13, Q-T), and they may be accompanied by 
one or two accessory teeth (Fig. 13, 0,P,S). Thoracic uncini 
possess 5-7 teeth in a single row (Fig. 13, U). Abdominal 
uncini are similar, with 5-6 teeth; anteriorly saw- and rasp- 
shaped uncini may occur in a single row, posteriorly all uncini 
are rasp-shaped. 

Live material. As observed in material collected in 1990, 
radioles are faintly yellow to lemon, operculum is transpar- 
ent, almost colourless. Thorax ventrally with 2-\ bright red to 
orange globules arranged in a V, presumably thoracic glands; 
body transparent with yellow tinge, brownish gut. 

Etymology. Named after Gayle Plaia who, when working 
at the Florida Marine Research Institute, first observed ITS 
in one of the species, S. ypsilon, from the Gulf of Mexico. 

Habitat and distribution. S. plaiae is a shallow water 
species occurring in coral reefs, and has hitherto been col- 
lected only from the type locality. 

Spiraserpula caribensis sp. nov. 

(Figs. 14, A-M; 15, A-Y; 16, A-K; 3, L; P1.4, E & F; 

P1.5,A-E) 

Material examined. 

Curasao (Neth. Ant.): 1. Awa Blancu, coral debris barrier, 
20-30 cm, legit H.A. ten Hove, 15. ix. 1975, Stn. 75-38 
(HOLOTYPE & 3 PARATYPES: ZMA V.Pol. 3715; 2 
PARATYPES USNM 130987; 4 PARATYPES each: AM 
W20157, NSMT, ZMK). 2. Awa Blancu, 3-4m, legit H. A. 
ten Hove, 14.x. 1975, Stn. 75-37 (1 specimen, HUJ). 3. Awa 
Blancu, coral debris, near Lagoen Blancu, 30-50 cm, legit 
H. A. ten Hove, 30.vii.1970, Stn. 2090 (several subsamples 
BM(NH) 1992.25-31, FSBC I 39195, ZMA V. Pol. 3716, 
ZMB). 4. Lagoen Blancu, coral debris barrier, Halimeda, 
20-30 cm, legit H. A. ten Hove, 15. ix. 1975, Stn. 75-36 (2 
out of several specimens, RMNH 18174). 5. Awa di 
Oostpunt, coral debris barrier, 30-50 cm, legit H. A. ten 
Hove, 3.x. 1975, Stn. 75-77 (1 out of few specimens, 
BM(NH) 1992.10-11. 6. St. Jorisbaai, Peninsula Groot St. 



Fig. 13 Spiraserpula plaiae sp. nov. A, Aggregation of tubes, showing fine transverse growth markings, granular overlay in places. B, 
Juvenile tube. C-E, Fragments of tubes showing serrated ventral ridge (C & D, paratype 3), (E, paratype 2). F, Holotype showing variant 
form of ventral ridge with isolated teeth, and ventro-lateral ridge. G-I, Paratype 4: G, posterior end of tube and worm. H, Portion of tube 
showing ventral ridge. I, Dorsal ridge on columella-shaped axis. J-L, Two views of paratype 4, and its operculum: J, (Operculum not seen) 
ventral longitudinal abdominal groove, and dorsal groove within 2nd coil; K, showing operculum; L, Another view of operculum. M, 
Paratype 2. N & O, Bayonet chaetae from holotype. P-T, Bayonet chaetae from paratype. U, Thoracic uncini from holotype. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



69 




70 



T.G. PILLAI AND HA. TEN HOVE 




Fig. 14 Spiraserpula caribensis sp. nov. A-J, From Florida, Stock Island, Stn. 7B. A, Tube with granular overlay and longitudinal ridges. 
B-F, ITS seen in various tube fragments: B, bottom cross-section showing median dorsal ridge on convex wall, narrow lateral ridge on 
either side, and serrated ventral ridge (barely visible); middle cross-section with serrated ventral ridge along concave wall; C, ventral ridge 
in sectional view (barely visible), and dorsal and lateral ridges; D, substratal view of first formed coil of tube opened to show smooth dorsal 
and serrated ventral ridge; E, smooth dorsal ridge in the first formed coil, on the right; F, uncoiled part of tube showing smooth dorsal 
ridge tapering at both ends. G, Anterior end of specimen, with thoracic glands, and showing dorsal and ventral abdominal grooves. H, 
Radioles of same specimen with left and right filamentous rudimentary opercula. I & J, anterior end of a worm with two views of collar. 
K-M, Bayonet chaetae. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



71 



Joris, muddy pebbles, Thalassia flat, few corals, from 
limestone boulders, 30 cm, legit H. A. ten Hove, 
10. ix. 1970, Stn.2096 (2 out of several specimens, BM(NH) 
1992.20-24, RMNH 18175). 7. St. Jorisbaai, Koraal Tabak, 
Punta Blanco, undersides of boulders, on rocky debris, 
20-30 cm, legit H. A. ten Hove, 15. xi. 1988, 9. i. 1990, near 
Stn. 75-30 (5 out of several specimens, MCZ, ZMH). 8. St. 
Jorisbaai, entrance channel, W, boulders and large metal 
poles in surf; from undersides crusts of Spiraserpula, legit 
H.A. ten Hove, 16. i. 1990 (clusters, AM W20341, HUJ). 
Aruba: 9. Spaans Lagoen, SE of bridge, rocks, etc., at 
floodgate, mud, Rhizophora, 0-2.0 m, legit P. Wagenaar 
Hummelinck, 24.iii.1970, Stn. 1673 (2 specimens BM(NH) 
1992.12-13). 10. Andicuri, cape W of beach, windward 
side, rockpool, exuberant coral growth, strong wave 
action, 0.5 m, legit H. A. ten Hove, 20.viii. 1970, 
Stn.2034B (several fragments of tubes, 2 incomplete 
worms, ZMA V. Pol. 3719). 

Barbuda: 11. Great Lagoon, Lobster Point, N. of Palm 
Beach, Thalassia and Halophila, 0-1.0 m, legit P. 
Wagenaar Hummelinck, 23.vii.1967, Stn. 1534 (3 out of 
several specimens, ZMA V. Pol. 3725). 
Bonaire (Neth. Ant.): 12. Lac, dam, beachrock in current 
behind surf, 5-10 cm, from crevices in beachrock, legit H. 
A. ten Hove, 15.vii.1970, Stn. 2123 (1 specimen, USNM 
130986). 13. Lagun, N shore, 500 m from entrance, rock, 
boulders, 0-50 cm, from undersides of boulders, legit H. 
A. ten Hove, 23. vi. 1970, Stn. 2129 (3 out of several 
specimens, ZMAV. Pol. 3720). 14. Bonaire, Karpata, reef, 
10 m, cryptic, legit H. A. ten Hove, 9.xi.88 (1 tube, 
BM(NH) 1992.14). 

Jamaica: 15. Drunkeman's Key, sandy debris, 0-0.5 m, 
legit P. Wagenaar Hummelinck, 15. vi. 1973, Stn. 1683, 
(ZMA V. Pol. 3723). 

Puerto Rico: 16. La Parguera, E, glade in mangroves, 
Thalassia beds, muddy sand, from between boulders, 
20-30 cm, legit H. A. ten Hove, 1.x. 1970, Stn. 2135 (3 
specimens, ZMA V.Pol. 3724). 

Panama: 17. Gatun Locks, walls of outer platform, lower 
W chamber. Pan. Survey, 20.iii.1972, Pacific Stn. 81-1, M. 
L. Jones coll., USNM No. 58661 (2 specimens without their 
tubes). 18. Same, Stn. 81-2, M. L.Jones coll., USNM 
No. 58662, (1 specimen with its tube). 

Florida: 19. Safe Harbour, Stock Island (near Key West), 
Florida Keys, 5 m, from chunks of calcareous materials 
(shells, barnacles, etc.) cemented together and covered 
with serpulids and small cirratulids, legit R. Chesher and 
C. Hamlin, 17.vii.1970 and l.vi.1971, Stn.7B, (22 out of 
several specimens, USNM 130988, BM(NH) 1992. 15-19, 
ZMAV. Pol. 3721 (10+ specimens from l.vi.1971). 20. Off 
Egmont Key, 27. 0°37.0'N, 83°01.5'W, sea buoy, 18 m, 
scarce sponges and corals, 2 cm of soft sludge on lime- 
stone, many serpulids, legit H.A. ten Hove and T. Perkins, 
2.U980, Stn. EJ. 80002, (9 out of several specimens, ZMA 
V. Pol. 3722, FSBC I 39202). 

Type locality. Curasao (Netherlands Antilles). 

Description. 

tubes: Light to bright pink or rose coloured. They form 
mutually bonded aggregations of a few to several individuals. 
Their external diameter is generally about 1.0 mm, maxi- 
mally 1.5 mm. There are three longitudinal ridges, one 
median and one along each lateral margin, which may be 



indistinctly developed in places (Figs. 14, A; 15, A). Narrow 
transverse ridges may be developed to various extents 
(Fig. 15, A). Some of the tubes end anteriorly in 4 rounded, 
anteriorly-directed lobes. A transparent to translucent granu- 
lar overlay is present. The granulations are larger and more 
densely laid along the ridges. The pink colour is faint along 
the longitudinal ridges, as seen through the transparent 
granules, but form of a pair of bright longitudinal bands 
between the ridges. Branching tubes, difficult to observe 
since they form dense aggregations, have been observed in 
material from Curacao (Stn. 2090, 2096, 75-38), and from 
Bonaire (Stn. 2123). 

ITS consist of a serrated ventral ridge along the concave 
wall (Figs. 14, B,D; 15, P), and a smooth dorsal ridge 
opposite (Figs. 14, B, D-F; 15, 0,P). The dorsal ridge is 
nearly tongue-shaped in cross-section, with a gradual 
decrease of its height, thickness and width of the widest part 
both anteriorly and posteriorly. This is occasionally more 
clearly seen in the non-coiled portions of tubes (Fig. 14, F). 
The dorsal ridge may be situated on a columella-shaped axis 
in tubes coiled upon themselves (Fig. 14, E). They usually 
also possess a short accessory dorso-lateral ridge on either 
side of the dorsal ridge (Figs. 14, B & C; 3, L). The inside of 
the tube may have a light caramel to light brown lining. The 
mid-ventral and mid-dorsal longitudinal abdominal grooves 
of the worm are applied to the serrated ventral and smooth 
dorsal ridges, respectively. 

worms: The longest available complete worm is from 
Florida. It has a total length of 12.8 mm, thoracic width of 
0.5 mm, abdominal length of 9.7 mm, and has 91 segments, 
with capillaries commencing on the 80th. There are four 
radioles and a rudimentary operculum on each side. Fully 
developed opercula are absent in all the specimens, being 
represented by a long and filamentous rudimentary opercu- 
lum on each side (Fig. 14, H). The highest number of radioles 
is 6 pairs, the longest measure about 2.1 mm, and end in 
slender pinnule-free tips which are 1/5-1/6 their entire length 
(Fig. 14, H). Radioles have up to 12 pairs of pinnules each, as 
could be observed in living material. The smallest worm is a 
juvenile from Curaco (Stn. 75-77) which has a total length of 
3.7 mm, a thoracic width of 0.45 mm, abdominal length of 
2.0 mm, and has 20 segments, with capillaries in the last 5. It 
has 4 radioles on the left and 5 on the right, in addition to the 
rudimentary opercula. 

Two reddish to reddish-brown clusters of prostomial ocelli 
are present. The median lobe of the collar is sub-rectangular, 
with rounded lateral borders and a smooth medial notch 
(Fig. 14, I & J). Five to seven globular ventral thoracic glands 
are present (Fig. 14, G), more or less arranged in a V. 
Whether they are responsible for secreting the brownish 
inner lining of the tube or not has to be further investigated. 

A summary of data is presented in Table 13. Similar data 
from the Florida material are provided in Table 14. 

The bayonet collar chaetae, which number 3 or 4 fully 
formed ones per fascicle and, usually, a developing one deep 
within, are unique among the species of Spiraserpula and of 
Serpula that have hitherto been described. Their blades are 
conspicuously short, unserrated and dagger-shaped (Figs. 14, 
K-M; 15, B-I, Q-W; P1.4, E & F). The number of large 
conical teeth on the basal boss is usually 3 or 4. Often there 
are 2 large teeth with 1 or 2 smaller ones in between (Fig. 14, 
K-L; 15, B-I). In the specimens from Gatun Locks, Panama, 
the number of teeth is usually 4 or 5 (Fig. 15, Q-W). These 
dagger-shaped bayonet chaetae were noted and figured in the 



72 



T.G. PILLAI AND H.A. TEN HOVE 



Table 13 5. caribensis sp. nov. A summary of data from four 
samples from Curacao (Stns. 75-38, 75-36 and 75-77 and 2096). 



No. of specimens (n=10) 
No of radioles per side 

No. of specimens (n = 14) 
No. of thoracic chaetal tufts 

No. of specimens (n = 10) 
Thoracic membrane ends 



2 7 1 
6/5 5/5 5/4 

2 13 3 4 1 

9/8 9/7 8/8 8/7 7/7 7/6 

12 4 2 1 

5/4 5/3 4/4 4/3 3/3 



Table 14 5. caribensis sp. nov. A summary of data from the 
Florida material. 



No of specimens (n=ll) 
No of radioles per side 

No. of specim.(n=29) 
No. of trior, chaet. 

No. of specimens (n=26) 
Thor. membranes end on 



2 
6/6 



5 
5/5 



4 

4/4 



18 13 6 12 2 4 1 

10/6 9/8 9/7 8/8 8/7 8/6 7/7 7/6 7/5 6/6 

3 3 4 9 7 

5/4 5/3 4/4 4/3 3/3 



unpublished research of M. van Vliet and R. Fijn (see 
acknowledgements) . 

The blades of developing bayonet chaetae deep within the 
fascicle are similar to the fully formed dagger-shaped bayonet 
chaetae, indicating that the latter have not resulted from wear 
and tear of bayonets with tapered tips. Occasionally, a 
developing chaeta with a truncated blade and tapered tip 
(Fig. 15, F,S), occurs deep within a fascicle, which provides a 
clue to the origin of the former. Reduction in length of the 
blade together with extension of the unserrated notch has 
resulted in stout, truncated bayonet chaetae, with smooth 
and dagger-shaped blades. 

Thoracic uncini (Fig. 15, J) usually possess 6 teeth, and 
anterior abdominal uncini (Fig. 15, K,X) 4 or 5, in a single 
row. Posterior abdominal uncini are rasp-shaped (P1.5, A). 
Flat trumpet chaetae number up to about 5 in each bundle, 
and their triangular distal ends bear a hook-shaped process 
on one side, and the other side is drawn out into an acute 
angle (Fig. 15, L-N, Y; P1.5, B). 

Collections from other localities. The specimens from 
the other localities listed above agree with those from the 
type locality. However, the smaller size of the tube and 
chaetae of the specimens from Gatun Locks, Panama, and 
the highly branched tubes of the specimens from Grenada, 
are worth noting. 

Live material. As observed in material from Curacao, 
radioles are colourless, transparent to transparently orange, 
sometimes with reddish pinnules. Base of branchial lobes and 
the collar may be tinged with purple. Branchial eyes not 



present. Body predominantly transparent orange, thorax 
ventrally reddish. 

Etymology. The name acknowledges the fact that this 
appears to be the most widely distributed species of Spiraser- 
pula in the Caribbean. 

Habitat and distribution. S. caribensis inhabits shallow 
water, intertidally down to a few metres in the Caribbean, to 
18 m in the E. Gulf of Mexico (temperature submerged ?). It 
occurs in a variety of habitats, from rockpools to the under- 
sides of boulders in mangrove glades. It is able to survive well 
in somewhat muddy environments, always, however, cryptic 
between piles of rock or similar hard substrata. 

It appears to be widely distributed in the Caribbean and 
Gulf of Mexico, from Florida to Barbuda and Panama. 

A population from Grenada, with frequently branching 
tubes and which is, for the present, regarded as belonging to 
5. caribensis, is described below (Fig. 16, A-K): 

Material examined. 

Grenada (Caribbean), Hog Island, near Pt. Salines, 0-1.5 m, 
Rhizophora, mud, legit P. Wagenaar Hummelinck, 
8.vii.l967, Stn. 1550 (5 specimens and 4 tubes, ZMA V.Pol. 
3706, USNM 130985, BM(NH) 1992.32). 

Description. 

tubes: Dark pink to rose coloured. Except for their posterior 
ends, they are all uncoiled, conspicuously branched, and 
attached to the substratum throughout (Fig. 16, A). A granu- 
lar overlay is present, larger granules constituting a median 
longitudinal ridge and a pair of lateral ridges (Fig. 16, A,B). 
The colouration is darker pink between the median and 
lateral ridges. Fine transverse ridges may be present in 
places. The lumen of the tube is continuous with that of the 
branches. 

ITS are similar to those of S. caribensis. However, the 
tongue-shaped cross-section of the dorsal ridge is somewhat 
more pronounced. 

worms: Three worms were taken out of the tubes, of which 
the longest (Fig. 16, C,D), has a total length of 9.2 mm. There 
are up to 5 pairs of radioles and a rudimentary operculum on 
each side. The radioles are up to about 2.0 mm long, 1/6-1/8 
of which constitute pinnule-free tips. Measurements and 
meristic data are given in Table 15: 

Two clusters of prostomial ocelli are present. All three 
specimens bear 7 thoracic chaetal tufts on the left and 6 on 
the right. The thoracic membranes end on the third thoracic 
chaetiger on both sides in the first specimen, but are damaged 
in the others. A pair of ventral thoracic glands is present 
(Fig.l6,C). 

Each bayonet chaeta typically consists of a short, serrated 
blade, and an unserrated notch and a tapered tip (Fig. 16, 
E-K). There are 2 or 3 large teeth on the basal boss, and a 
few accessory teeth. Older chaetae in a fascicle which have 



Fig. 15 Spiraserpula caribensis sp. nov. A-E & J-N, From Curacao, St. Jorisbaai, Stn. 2096. F-I, from Curacao, Lagoen Blancu, Stn. 75-36. 
O-Y, From Panama, Gatun Locks: (O-S, from Stn. 81.1; T-Y, from Stn. 81.2). A, Tubes showing granular overlay, external ridges and 
transverse wrinkles. B-E, Bayonet chaetae from the same fascicle with short dagger-shaped blades. F-I, Bayonet chaetae from fascicle of 
another specimen: F, Newly formed, deep within fascicle; G-I, Older chaetae. J, Thoracic uncini. K, Anterior abdominal uncini, and L, 
flat trumpet chaetae, from same segment. M & N, Flat abdominal trumpet chaetae from other specimens. O & P, Tube opened 
substratally, viewed from two different angles, with worm in situ showing thoracic glands; O, with dorsal ridge only, and P, with both dorsal 
and ventral ridges. Q-S and T-W, Bayonets from two different fascicles. Note much smaller size compared with those of Florida (Fig. 14, 
K-L) and Curasao (Fig. 15, B-I) material', although drawn under same magnification. X & Y, Anterior abdominal uncini and flat trumpet 
chaetae from same segment. Note much smaller size than in Curasao material (Fig. 15, L-N). • 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 




74 



T.G. PILLAI AND HA. TEN HOVE 




Fig. 16 Spiraserpula caribensis sp. nov. A-K, From Grenada. A, Branched tube with granular overlay and longitudinal ridges. B, Branching 
point marked Y in A, magnified. C & D, Two views of a worm showing rudimentary opercula, thoracic glands (C), and dorsal and ventral 
longitudinal abdominal grooves. E-H, Four bayonets from a small specimen: A, Older chaeta with worn out tip; F-G, Chaetae with intact 
tips. I-J, Bayonets from a larger specimen: the oldest (K) with a worn out tip, and the other two with intact tips. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



75 




Fig. 17 Spiraserpula nudicrista sp. nov. From Bonaire. A, Tube, with granular overlay and longitudinal ridges. B-D, Tube fragments 
showing unserrated ventral ridge along concave wall and lack of a ridge on the opposite side. E-F Other tube fragments showing a narrow 
ventral ridge anteriorly, and a smooth and rounded edge to it posteriorly. J, First formed coil, lacking a dorsal ridge. K-L, Two different 
views of holotype showing prostomial ocellar clusters, as seen through the collar, the ventral longitudinal groove of the abdomen, and 
extent of thoracic membrane on the left side (K); L, Radioles with a pair of short club-shaped rudimentary opercula and moderately long 
pinnule-free tips. M & N, Paratype: M, tube with granular overlay, prostominial ocellar cluster of one side, and longitudinal abdominal 
grooves; N, branchial crown with a pair of short club-shaped rudimentary opercula, and long pinnule-free tips. 



76 



T.G. PILLAI AND H.A. TEN HOVE 



Table 15 S. caribensis from Grenada. Measurements and counts. 



Radioles 



Abdomen 



Total Thoracic Number 

Specimen length width Length Length of Capillaries 

no. (mm) (mm) (mm) No. (mm) segments on 



1 


9.2 


0.5 


2.0 5/5 6.6 


50 


44 


2 


8.1 


0.5 


1.2 4/3 4.7 


30 


7 


3 


3.1 


0.35 


0.6 3/3 1.8 


35 


10 



lost their tapered tips through abrasion may appear some- 
what like the bayonets of S. caribensis from elsewhere 
(Fig. 16, E, K; PI. 5, C-E), but the newly formed bayonets, 
within the fascicle, possess tapered tips. 

Habitat and distribution. Appears to inhabit shallow 
water and capable of withstanding the silty conditions found 
in mangrove backwaters. It was found on the inside of a dead 
oyster shell covered with much silt. 

Remarks. The extensively branching tubes and differences 
in the collar chaetae initially led us to consider the Grenada 
material as possibly belonging to a distinct species. However, 
branching as such, although inconspicuous, was also subse- 
quently observed in some specimens of S. caribensis from 
Bonaire (Stn. 2123) and Curacao (Stns. 2090, 2096,75-38; see 
above), in S. paraypsilon from Curacao (lO.i.90). Moreover, 
S. snellii, described later in this paper, revealed a schizont 
with parent in one tube. By itself, therefore, branching 
cannot be a good character to separate the Grenada material 
as a distinct species. The fully formed bayonet chaetae, 
including those within the fascicle, of S. caribensis proper, 
have short dagger-shaped blades with blunt tips, while blades 
of the Grenada material typically end in tapered tips. 
Although the tip of a fully formed chaeta in the Grenada 
material might be lost through abrasion (Fig. 16, E-K), those 
deep within the fascicle are tapered. 

Further work on additional material is necessary to deter- 
mine whether frequent branching of the tubes and the 
features of the bayonet chaetae are consistent, and whether 
there are other characters which would justify the separation 
of the Grenada material into a distinct species or not. 

Spiraserpula nudicrista sp. nov. 

(Figs. 17, A-N; 18, A-O; 3, F; PI. 3, A-D) 

Material examined. 

Bonaire (Neth. Ant.): 1. Karpata, reef, cryptic, 10 m, legit H. 

A.ten Hove, 9.xi.l988, (HOLOTYPE & PARATYPE: 

ZMAV.Pol.3711). 

Curacao (Neth. Ant.): 2. Savonet, E of Boca Braun, reef, no 

sand, about 22 m, from corals, some dead, legit H. A. ten 

Hove, 28.xi.1970, Stn. 2101 (PARATYPES 2 & 3: BM(NH) 

1992.61 & 62). 

Type locality. Bonaire (Netherlands Antilles). 



Description. 

tubes: White to creamish white and have a conspicuous 
granular overlay (Fig. 17, A,E,M). They may be covered over 
by encrusting calcareous organisms. They are trapezoidal in 
cross-section, with two longitudinal ridges along the crest of 
the tube and one along each flank (Fig. 17, A). The maximum 
external tube diameter of the holotype is 1.0 mm. 

ITS consist of an unserrated ventral ridge which is rounded 
and smooth towards its middle (Figs. 17, H; 3, F), from where 
it decreases in thickness and height both anteriorly and 
posteriorly (Fig. 17, C-E, F-G). A dorsal ridge is generally 
absent, even on the convex pulley-shaped posterior end 
(Fig. 17, I). However, paratype 1 from Bonaire showed some 
isolated dorsal teeth. The mid-ventral longitudinal groove of 
the abdomen (Fig. 17, M) is applied to the unserrated ventral 
ridge . 

worms: Only two worms were yielded by the tubes from 
Bonaire. The complete holotype has a total length of 
15.6 mm, a thoracic width 0.7 mm, an abdominal length 
11.9 mm and about 101 segments, with capillaries on the last 
7. The radioles are 2.5 mm in length, and their pinnule-free 
tips of 0.6 mm are comparatively long (Fig. 17, L, N). The 
paratype is incomplete posteriorly. 

The holotype has 9 pairs of radioles while the paratype has 
8 pairs. Both specimens have a short filamentous rudimentary 
operculum on each side (Fig. 17, L). Two clusters of prosto- 
mial ocelli are present, and are seen as conspicuous brown 
patches through the collar (Fig. 17, J,K,M). This is in contrast 
to the other known members of the genus in which they can 
be seen when viewed from the anterior end with the radioles 
removed or when mounted. 

Both specimens have 8 pairs of thoracic chaetal tufts, and 
the thoracic membranes end on the fourth chaetiger on the 
left (Fig. 17, K) and the 5th on the right. Ventral thoracic 
glands appeared to be absent. 

The collar fascicles of the holotype possess four bayonet 
chaetae with long serrated to pilose blades and several conical 
teeth on the basal boss (Fig. 18, A-D; PI. 3, A). There may be 
a number of accessory teeth arranged around the bases of the 
larger teeth, which are lacking in the paratype (Fig. 18, E-G). 
The unserrated notch is short. Thoracic uncini (Fig. 18, H; 
PI. 3, B) and anterior abdominal uncini (Fig. 18, I; PI. 3, C) 
possess 4 or 5 teeth arranged in a single row. There are about 
4 flat trumpet chaetae in each abdominal fascicle (Fig. 18, 
J;P1.3, D). An anterior hook, as in most of the species of the 
group, cannot be discerned, all distal teeth appearing more or 
less equally developed. 

Tubes of the specimens from Curacao are similar to those 
from Bonaire with regard to colour, form and ITS (Fig. 18, 
K). Their maximum external diameters are 1.1-1.2 mm. Two 
of them yielded worms which are incomplete posteriorly. 
Some data from them are presented in Table 16. 

Both specimens possess a rudimentary operculum on each 
side (Fig. 18, L,M). Bayonet collar chaetae (Fig. 18, N,0), are 
similar to those of the specimens from Bonaire, although 
their basal bosses are somewhat stouter. 

Remarks. A small fragment from the inside of the coil of 



Fig. 18 Spiraserpula nudicrista sp. nov. A-J, From Bonaire. A-D, Bayonet chaetae, holotype. E-G, Three, out of five, bayonet chaetae 
from paratype. H-J, from paratype: H, Thoracic uncini; I & J, flat trumpet-shaped chaetae and uncini from anterior abdomen. K-O, From 
Curaao. K, Tube, with granular overlay, external longitudinal ridges, and internal ventral longitudinal ridge seen through fractured end. L, 
Branchial crown of older specimen with pair of rudimentary opercula. M, Branchial crown of younger specimen, with pair of shorter 
rudimentary opercula. N & O, Two, out of four, bayonet chaetae. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



77 




78 



T.G. PILLAI AND H.A. TEN HOVE 



Table 16 S. nudicrista sp. nov. Some data on two specimens from 
Curacao. 





Specimen 1 


Specimen 2 


Width of thorax 


0.5 mm 


0.6 mm 


Length of radioles 


2.1 mm 


2.6 mm 


Pinnule-free tips (Figs.l7L,M) 


short 


short 


No. of radioles (L/R) 


7/8 


9/8 


No. of thoracic chaetal tufts (L/R) 


9/7 


8/6 


Thoracic membrane ends 


?/? 


4/4 


Length of abdomen 


? 


9.7 mm 


No. of abdominal segments 


? 


about 60 



paratype 1 showed a slightly concave to asymmetrical cross- 
sectional edge to the ventral ridge, and a few isolated teeth in 
the location of the dorsal ridge, somewhat similar to the 
condition in S. paraypsilon. There is also some similarity in 
the collar chaetae. 

ETYMOLOGY, nudus (L) = unadorned; crista = crest, ridge. 

Habitat and distribution. S. nudicrista is a shallow water 
cryptic species inhabiting coral reefs. It has hitherto been 
collected from Bonaire and Curacao. 

Spiraserpula sp. 
(Fig.19.A-C) 

Material examined. 

Curasao (Neth. Ant.): Piscadera Baai, outer bay in front of 

CARMABI, muddy reef, many sand spots, about 40 m, from 

dead corals, legit H.A. ten Hove, 9.vi.l970, Stn. 2054B (3 

empty tubes, and some abdominal fragments, ZMA V. Pol. 

3883). 

Description. Tubes are white, circular in cross-section. An 



erect portion shows a granular overlay, and an encrusting 
sponge at its base (Fig. 18, A). ITS characteristic of this genus 
are present in the coiled parts, and consist of an unserrated 
dorsal ridge and a serrated ventral ridge. The dorsal ridge is 
transparent, somewhat high, and has a smooth, somewhat 
T-shaped edge; it is spiral, on a columella-shaped axis in the 
spiral proximal portions of the tube (Fig. 18, B,C). The 
available portions of the worms were inadequate to assign the 
material to any of the other Caribbean species or a new 
species. 

Spiraserpula vasseuri sp. nov. 
(Figs. 20, A-H;21, A-K; 3, J) 

Helmut Zibrowius requested (pers. comm.) that the material 
on which he based his preliminary description of the present 
species in an unpublished manuscript be examined, and that 
it be included in this paper if it belonged to the present group. 
ITS are indeed present in this species, and its description 
follows: 

Material examined. 
Europa Island (Mozambique Channel): North Reef, Gabriel 
Cove grotto, 55 m, on oysters, legit Pierre Vasseur, scuba 
diving, 28.xii.1965 (HOLOTYPE: USNM 46475, 6 
PARATYPES USNM 46476). 

Type locality. Europa Island. 

Description. 

tubes: The colour is mostly whitish, with a very faint pinkish 
to orangish tinge seen in places at certain angles of illumina- 
tion. They are sinuous, coiled and bonded together, espe- 
cially at their bases. A granular overlay is present. The 
anterior portions are squarish to trapezoidal in cross-section 
(Fig. 20, A-C). The dorso-lateral angles may be incompletely 
developed in places and represented by a pair of incomplete 
longitudinal ridges; an additional incomplete ridge may be 





. 5mm 



Fig. 19 Spiraserpula sp. A-C, Three specimens from Curacao. A, Erect part of tube with encrusting sponge on its base. B, A tube opened to 
show spiral dorsal ridge in its first formed coil. C, Aggregation of tubes opened to show variations of dorsal ridge. 






ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



79 




. 5mm 





. 5mm 





. 5mm 



P 



. 5mm 



Fig. 20 Spiraserpula vasseuri sp. nov. A-H, paratypes. A-C, Anterior tube fragments showing granulations, transverse ridges and roughly 
trapezoidal external outline. A, Shows incompletely formed external longitudinal ridges, and a thickened peristome towards the posterior 
end of B. D & E, Two views of same posterior coil showing the unserrated dorsal ridge in both, and a serrated ventral ridge consisting of 
separate teeth in D. F & G, Two views of same anterior tube fragment with an attached posterior coil opened (on the right). The anterior 
tube fragment has two peristomes; the posterior coiled part (G)has been opened to expose the unserrated dorsal ridge and serrated ventral 
ridge (G). H, Thorax showing prostomial ocelli, collar, thoracic membranes and chaetigers. 



80 



T.G. PILLAI AND HA. TEN HOVE 



present along each flank (Fig. 20, A). Transverse ridges are 
present, which may be thickened in places, representing 
peristomes (Fig. 20, B,F,G). Although broken into fragments 
during collection, total lengths appear to have been between 
30-40 mm, and their maximum external diameters up to 
about 3.0 mm. Their fractured ends show two concentric 
layers of different consistency and thickness, an inner one 
that is more vitreous and transparent than the outer which is 
white and opaque. Their lateral margins are fragile and 
chambered, with thin walls. 

The posterior ends of the tubes are coiled. ITS consist of a 
low unserrated dorsal ridge, and a serrated ventral ridge 
(Fig. 20, D-G). The latter may be represented by a row of 
separate teeth (Figs. 20, D; 3, J). 

worms: The total length of the worms, based on the frag- 
ments, exceeds 15.0 mm. The branchial crown is 4.0-5.0 mm 
long, and each side bears 8-10 radioles and an operculum or a 
rudimentary operculum. The opercular peduncle is long and 
slender, of the same thickness as the radioles. One of the 
specimens has a well-developed operculum on one side, and 
another, much smaller, but similar operculum on the other. 
The operculum is big and short, massive, bell-shaped, and 
slightly concave distally. The radii end in large, rounded 
marginal lobes, and range from 10 to 15 in number (10 in 1, 
11 in 3, 12 in 1, and 15 in 1). The number of thoracic segments 
per side varies from 10 to 14. A pair of small ocellar clusters is 
present. Collar large, roughly divided into three large ventral 
lobes and a pair of latero-dorsal lobes. Thoracic membranes 
are broad up to the third segment, after which they narrow, 
and do not form an apron. 

The longest opercular peduncle (holotype) together with 
its operculum is 5.5 mm long. The operculum (Fig. 21, 
A,C,D) is separated from the peduncle by a faint constric- 
tion, where the peduncle is only 1/2-1/3 the diameter of the 
base of the operculum. The variations in the dimensions of 
opercula of the older specimens are as follows: length: 
0.6-0.7 mm; width: 0.55-0.6 mm. They are bell-shaped, with 
a small shallow concavity distally. They have a thick and 
transparent cuticle (Fig. 21, A,C,D). The second radiole of 
the opposite side is modified into a rudimentary operculum 
(Fig. 21, B). The radioles end in short pinnule-free tips which 
are about 1/10— l/15th the total length of the radioles (Fig. 21, 
A-C). 

One of the specimens, a juvenile, provides an indication of 
the possible ontogenetic changes in the operculum of this 
species. Unlike in the adults, where peduncle and operculum 
are markedly separated from each other, the slender 
peduncle of the juvenile merges gradually into the base of the 
operculum. In addition, the shape of the latter is an elongated 
funnel, and its distal end is convex (Fig. 21, E). 

The collar fascicles may bear up to about 5 fully formed 
bayonet chaetae and one developing deep within. Each 
possesses a long serrated blade, a short unserrated notch, and 
several moderately large teeth on the basal boss (Fig. 21, 
F-K). Thoracic uncini show 5-6 teeth in side view; however, 
in oblique edge view it is evident that they are saw-rasp 
shaped, with an anterior single row and a posterior cluster of 
teeth (Fig. 21, L). This is more clearly seen in the anterior 



abdominal uncini (Fig. 21, M). In side view, the number of 
teeth in the latter vary from 4 or 5 towards the lateral end of 
the torus to 7 at the dorsal end. Flat trumpet chaetae number 
9-11 per bundle. Their distal ends terminate in a slender 
hook-shaped process on one side and are drawn out into an 
acute angle on the other (Fig. 21, N). 

Etymology. As suggested by Zibrowius (pers. comm.), the 
species is named after its collector, P. Vasseur. 

Habitat and distribution. A reef dweller found on oyster 
shells in submarine caves at depths of around 55 m. Hitherto 
collected only from the Mozambique Channel. 

Spiraserpula deltoides sp. nov. 

(Figs. 22, A-N; 3, C) 

Material examined. 

Lesser Sunda Islands, Sumba (Indonesia): Snellius II 4.051, 
NE coast of Sumba, E. of Melolo 09°53.5'S 120°42.7'E, 
75-90 m. (HOLOTYPE & 1 PARATYPE (empty tube): 
RMNH 18296; 3 PARATYPES: ZMA V. Pol. 3736; 
2 PARATYPES: BM(NH) 1992.37 & 38). 

Type locality. Sumba (Indonesia). 

Description. 

TUBES: White, small, and spirally coiled upon themselves. 
They are squarish in cross-section, smooth and rounded 
dorso-laterally, and with a shallow longitudinal depression in 
between (Fig. 22, A). They have an extremely fine granular 
overlay, which can only be seen at certain angles of illumina- 
tion, and very fine transverse grooves. The coil diameter is 
generally about 3 mm, maximally 9 mm; the maximum exter- 
nal tube diameter is generally 0.7 mm, maximally 1.3 mm. In 
two of the tubes an inner transparent lining was observed. 

ITS consist only of a serrated dorsal ridge along the convex 
wall of the tube (Figs. 22, B,C; 3, C). The serrations are 
delta-shaped, mostly separate, and opaquely white in colour. 
worms: The holotype (Fig. 22, D) is 5.0 mm long, 0.35 mm 
wide in the thorax and its abdomen is 3.2 mm long. One 
paratype is incomplete posteriorly, the other is 8.0 mm long, 
with an abdomen of 3.5 mm. Some measurements and counts 
are given Table 17: 

The operculum is bell-shaped, with a shallow distal concav- 
ity extending inwards as far as the inter-radial grooves. The 
radii end in rounded marginal lobes, the constriction between 
operculum and peduncle is sharp, and the diameter of the 
distal end of the peduncle is about 1/2-3/4 that of the 
proximal part of the operculum (Fig. 22, D-G). The rudimen- 
tary operculum is 1.5 mm long, thread-shaped. The radioles 
end in short pinnule-free tips, about 1/7 the total length of the 
radioles. Two clusters of prostomial ocelli are present. It is 
difficult to determine whether ventral thoracic glands are 
present. Thoracic membranes do not extend to the end of the 
thorax, but exactly where they end cannot be located, it may 
be at the 7th chaetiger in one paratype. The abdomen of the 
holotype has about 67 segments, with capillaries on the last 8 
or 9; the complete paratype has 85 abdominal segments, 24 
with capillaries. The abdomen of the incomplete paratype is 



Fig. 21 Spiraserpula vasseuri sp. nov. A, Holotype. B-N, Paratypes. A, The left branchial crown and three views of the operculum and its 
slender peduncle. B, Left branchial crown and rudimentary operculum from another specimen. C & D, branchial crowns and differrent 
views of the opercula of two other specimens. E, Two views of the convex operculum of a juvenile. F-K, Bayonet collar chaetae bearing 
several teeth on the basal boss, and a short unserrated notch. L, Thoracic uncini, with more than one row of teeth towards their posterior 
ends. M, Anterior abdominal uncini. N, Bundle of anterior abdominal chaetae with flat trumpet-shaped ends. 






ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



81 




82 



T.G. PILLAI AND HA. TEN HOVE 



Table 17 5. deltoides sp. nov. Some measurements and counts on 
the holotype and 2 paratypes. 





Holotype 


Paratype 
1 


Paratype 
6 


Length of operculum and peduncle 

Length of operculum 

Diameter of operculum 

No of opercular lobes 

No. of radioles (L/R) 

No. of thoracic chaetal tufts (L/R) 


0.94 mm 
0.38 mm 
0.44 mm 

22 
5/6 

8/7 


1.2 mm 
0.52 mm 
0.35 mm 

26 

6/6 

6/8 


3.2 mm 
0.48 mm 
0.41 mm 

22 
7/7 
111 



12.5 mm long, has about 75 segments, and the latter bear 
reddish-brown granular material ventro-laterally. 

The collar fascicles bear 2-6 bayonet chaetae. Each has a 
long serrated blade, a short unserrated notch, and about 2-6 
teeth on the basal boss (Fig. 22, H-I, L-N). Thoracic uncini 
bear about 5 teeth in a single row (Fig. 22, J,K); anterior 
abdominal uncini are similar and bear 5-7 teeth. 

Etymology. The specific name refers to the delta-shaped 
serrations of the internal dorsal ridge. 

Habitat and distribution. Found on calcareous stones at 
depths of 75-90 m. Hitherto collected only from Sumba 
(Indonesia). 

Spiraserpula sumbensis sp. nov. 
(Figs. 23, A-U; 3, H) 

Material examined. 

Sumba (Indonesia): Snellius II 4.051, NE coast of Sumba, E 
of Melolo, 09°53.5'S 120°42.7'E, 75-90 m, (HOLOTYPE: 
RMNH 18297; 1 PARATYPE: ZMA V. Pol. 3737; 1 
PARATYPE: BM(NH) 1992.72). 

Type LOCALITY. Sumba (Indonesia). 

Description. 

tubes: White to very faintly pinkish. A small species with 
external tube diameter only up to about 0.5 mm, and a lumen 
of about 0.25 mm wide. A granular overlay consisting of 
extremely fine granules can be seen under special illumina- 
tion. Tubes are circular in cross-section and bear faint trans- 
verse wrinkles (Fig. 23, A-C, O). 

ITS consist of a dorsal ridge and a ventral ridge, which are 
both unserrated, wedge-shaped in cross-section (Fig. 3, H), 
and partially divide the lumen into somewhat asymmetrical 
left and right halves (Fig. 23, C,D). The two ridges are light 

Table 18 S. sumbensis sp. nov. Measurements and counts. 



Length of op. & peduncle (mm) 
Length of operculum (mm) 
No. of lobes 
No. of radioles 
No. of thoracic chaetal tufts 
Thoracic membrane ends 
(Fig.23,Q) 



Paratype 


Paratype 




1 


2 


Left side 


Right side 




1.2 


1.0 


1.2 


0.36 


0.36 


0.35 


19 


21 


17 


5 


5 


4/4 



pink and opaque. In cross-section they consist of a lens- 
shaped whitish kernel in the inner hyaline tube layer; the 
outer tube layer is opaque. 
worms: The holotype (Fig. 23, E-G), has a total length of 

7.0 mm, thoracic width of 0.26 mm, an abdominal length of 

5.1 mm and 66 segments, with capillaries on the last 17. The 
length of the operculum plus peduncle is 1.3 mm, the length 
and diameter of the operculum 0.38 mm and 0.26 mm, 
respectively. The operculum is zygomorphic (Fig. 23, E,F). It 
has a distal concavity which extends as far as the inter-radial 
grooves. The 15 radii end in somewhat acutely triangular 
marginal lobes with smooth tips. The peduncle is slender, but 
somewhat expanded before the constriction below the oper- 
culum. There are 5 radioles on each side, with the operculum 
on the left side and a short filamentous rudimentary opercu- 
lum on the right (Fig. 23, E). The short pinnule-free tips are 
about 1/7-1/8 the total length of the radioles. Thoracic 
chaetal tufts number 8 on each side. The thoracic membrane 
ends on the fifth chaetiger on the left, but it is difficult to 
determine its extent on the right. One tiny prostomial eye 
appears to be present on the right side, the left side is 
damaged. Thoracic glands could not be detected in the 
material. 

One paratype (Fig. 23, O-Q) has an incomplete abdomen. 
It is the first specimen encountered in this genus with two 
equally well-developed opercula (Fig. 23, 0,P). The thorax of 
the second paratype is missing, the remaining abdomen has 
54 segments, 12 of them with capillaries. Some measurements 
and other data are given in Table 18: 

The paratypes agree with the holotype with regard to the 
tube, operculum, radioles, and chaetae. The opercula are 
somewhat zygomorphic. 

Collar fascicles bear 4 fully formed bayonet chaetae in the 
holotype; 3 fully formed bayonet chaetae and a newly formed 
one deep within the fascicle in paratype 1. Each bayonet 
chaeta (Fig.23,H-K, R-U) consists of a long serrated blade, a 
moderately long unserrated notch, which is about 1/6-1/7 the 
length of the blade, and several teeth on the basal boss. 
Thoracic uncini appear to have a row of 7-9 teeth in side 
view, but more than one row as seen in edge view (Fig. 23, L). 
Anterior abdominal uncini are similar, but appear to have 
fewer teeth in side view (Fig. 23, M). Flat trumpets number 
four in each anterior bundle, their curved distal ends have a 
poorly developed hook on one side and are comparatively 
elongated on the other (Fig. 23, N). Capillaries occur in the 
posterior 12-17 chaetigers. 

Etymology. Named after the type locality. 

Habitat and distribution. Found on calcareous stones at 
depths of about 75-90 m. Hitherto collected only from 
Sumba (Indonesia). 

Spiraserpula iugoconvexa sp. nov. 
(Figs. 24, A-K; 25, A-Q; 3, 1) 

Material examined. 

NE Flores Sea to SW Banda Sea (Indonesia): 1. Taka Bone 

Rate(Tiger Islands), Snellius II 4.139B, S of Tarupa Kecil, 
06°30'S 121°8'E,depth -30 m, (HOLOTYPE: RMNH 18295; 
PARATYPE I: ZMA V.Pol.3735). 2.Tukang Besi Island, 
Binongko, Snellius II 4.044B, SW of Taipabu, Banda Sea, 
5°56'S 123°58.5'E, down to 25 m, (PARATYPE II: BM(NH) 
1992.39). Queensland (Australia): 3. Lizard Island, S. South 
Island, sloping silty reef, little coral cover, legit H. A. ten 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



83 




Fig. 22 Spiraserpula deltoides sp. nov. A-E & H-K, From holotype; F-G & L-N, from paratype. A, Tube viewed from above. B & C, Two 
views of same tube opened substratally to show deltoid dorsal ridge. D, Worm, showing operculum and dorsal abdominal groove. E, 
Anterior view of operculum. F & G, Two views of operculum. H & I, L-N, Bayonet chaetae. J, Thoracic uncini. K, Anterior abdominal 
uncini. L-N, Bayonet chaetae. 



Hove et al. 
W21676). 



Stn. 21,6.iii.l986 (1 specimen, tube; AM, 



Type locality. NE Flores Sea (Indonesia). 

Description. 

tubes: Bright rose, red in fresh material, with a translucent 
granular overlay, coiled posteriorly but not anteriorly. All 
three tubes of the type series were partially overgrown by 
encrusting bryozoans, making their surfaces irregular. At 
irregular intervals, there are also peculiar elongated struc- 



tures with a semilunar opening, which appear to be a hydro- 
zoan commensal, akin to Protulophila Rovereto, 1901 (vide 
Scrutton, 1975). These sparsely occurring structures are 
directed longitudinally or transversely as shown on the speci- 
men from from Stn. 4044B (bottom end of Fig.24, A). The 
anterior end of the tube from Tukang Besi Island is not 
attached to the substratum. Viewed dorsally, it is squarish to 
trapezoidal in cross section. It has two fairly distinct dorso- 
lateral ridges and a faint median one in places, transverse 
wrinkles which are occasionally thickened, and an expanded 



T.G. PILLAI AND HA. TEN HOVE 



peristome at its anterior end (Fig. 24, A). The ventral side of 
the unattached part, which commences from a swollen attach- 
ment to the substratum, does not show the granular overlay 
but only faint transverse wrinkles (Fig. 24, B). The peristome 
consists of a broad triangular dorsal lobe which is continuous 
with two narrow ventro-lateral lobes (Fig. 24, A,B).The 
inside of the tube is quite shiny. It attains a diameter of 
1.6 mm at the peristome, and 1.5 mm at the swollen com- 
mencement of the unattached part. 

ITS, which are present only in the posterior part of the tube, 
consist of an unserrated dorsal ridge (Figs. 24, C; 25, B,C), and a 
very short smooth ventral ridge (Fig. 3, 1), which is very short in 
the holotype (Fig. 25, D); in the specimen from Lizard Island the 
ventral ridge is smooth to scalloped. The dorsal ridge may be 
T-shaped in cross-section in places (Fig. 25, C) but appears 
irregular if damaged (Fig. 24, C, middle). In the middle region of 
the tube, tear-shaped depressions are present in the inner wall, 
up to 0.2 mm in size. 

worms: The holotype from Taka Bone Rate, broken in three 
parts (Fig. 25, E-G), has a total length of 31.5 mm, a thoracic 
width of 0.5 mm, an abdominal length of 27.0 mm and 117 
segments, with capillaries on the last 8. Its radioles are 
2.7-3.0 mm long, of which the slender pinnule-free tips consti- 
tute 0.3 mm. Paratype I lacks its branchial crown; it has a length 
of 5.0 mm, a thoracic width of 0.4 mm, an abdominal length of 
4.3 mm, and has 61 segments, with capillaries on the last 15 orl6. 
Paratype II, from Tukang Besi Island, lacks its radioles on the 
right side, and its abdomen is in several parts. Its thorax, 
however, is intact (Fig. 24, G,H). 

The operculum and peduncle measure 3.0 mm long in the 
holotype, 4.1 mm in paratype II. Other measurements and 
counts are given Table 19. 

The operculum is zygomorphic, and its distal end is quite 
different from that of other known species of the genus in being 
markedly convex (Figs. 24, D-F; 25, G-J). The cuticle is thick- 
ened and transparent, particularly in its convex distal end, the 
marginal lobes of the radii, and the asymmetrical projection at 
the base of the operculum. The number of radial lobes reaches 
about a dozen. There is a sharp constriction between the 
operculum and the peduncle, the latter being slender, except for 
a slight expansion before the constriction (Figs. 24, E,F; 25, J). A 
filamentous rudimentary operculum is present on the side 
opposite to that of the operculum (Fig. 25, G). 

The number of radioles per side reaches 14. Their short 
pinnule-free tips are about 1/7-1/8 the entire length of the 
radiole. Prostomial eyes were not found. Thoracic glands are 
present, transparent in the holotype and paratype II, light 
brown in paratype I. The number of thoracic segments per 
side is 7-8, and the thoracic membranes do not reach the last 
thoracic segment (Fig. 24, G-I). 

The abdomen of the holotype appears glandular ventrally, 
packed with eggs, and bears peculiar swellings (Fig. 25, E) which 
fit into corresponding depressions in the tube. It was not possible 
to find them in the damaged abdomen of paratype II, although 
this is a mature specimen too, and the inner tube wall shows 
tear-shaped depressions (0.32 x 0.22 mm); they are absent in 



Table 19 S. iugoconvexa sp. nov. Measurements and other data on 
type specimens. 



Holotype Paratype I Paratype II 



Length of operculum (mm) 


0.7 


? 


1.0 


Diameter of operculum (mm) 


0.5 


7 


0.7 


No. of radii 


12 


? 


11 


No. of radioles(L/R) 


10 


7/7 


14/? 


No. of thoracic chaetal tufts 


8/7 


111 


in 


Thoracic membrane ends 


111 


1/3 


5/4 



the juvenile paratype I. Possibly, these abdominal swellings are 
developed in older worms only. 

Collar fascicles of the holotype bear 4 bayonet chaetae 
each (Fig. 25, K-N). Each possesses a long serrated blade, a 
short unserrated notch and two teeth on the basal boss, one 
of which may be difficult to observe in side view since it lies 
directly behind the other. The number of teeth is clearly seen 
in one of the bayonets of paratype I which has its blade 
broken off at its base (Fig. 24, J), although it is difficult to 
observe in a newly formed chaeta from within the same 
fascicle (Fig. 24, K). Thoracic uncini bear 5 or 6 teeth (Fig. 25, 
O), and anterior abdominal uncini 4 or 5 teeth in a single row 
(Fig. 25, P). There are up to about a dozen flat trumpet 
chaetae in each bundle (Fig. 25, Q). Their distal ends bear a 
claw-shaped process on one side and are drawn out into an 
acute angle on the other. 

Etymology. Iugum (L) = yoke; convexus (L) = bulbous; 
refers to the zygomorphic, convex operculum. 

Material from other locality. The material from Liz- 
ard Island agrees closely with that of the type series with 
regard to collar chaetae, operculum and tear-shaped depres- 
sions in the inner tube wall. However, the ventral internal 
ridge has a scalloped edge, not smooth as in the Indonesian 
material. 

Habitat and distribution. A reef dweller occurring at 
depths of about 25 m. Hitherto collected from Indonesia 
(Flores Sea and Banda Sea) and Australia (Queensland). 

Spiraserpula snellii sp. nov. 

(Figs.26, A-X; 27, A-L; 28, A-V; 3, F) 

Material examined. 

Flores Sea, (Indonesia): 1. Taka Bone Rate (Tiger Island), 
Snellius II 4.139B, S. of Tarupa Kecil, 06°30'S 121°8'E, edge 
of reef flat, 30 m, (HOLOTYPE & 4 PARATYPES: RMNH 
18298; 4 PARATYPES (+ one abdomen & internal tube 
ridge): BM(NH) 1992.66-71; 5 PARATYPES & tube mate- 
rial: ZMA V. Pol. 3738; 3 PARATYPES & fragmentary tube 
material: USNM 130983 & 130984). 






Fig. 23 Spiraserpula sumbensis sp. nov. A-M, Holotype. O-U, Paratype. A-D, O, Tubes showing granular overlay and faint transverse 
growth ridges; A, An erect part; B, also showing body, operculum & radioles in situ; C & D, two views of same tube fragment showing 
wedge-shaped dorsal and ventral ridges, both unserrated. E, Operculum. F & G, Holotype. F, Radioles showing zygomorph operculum on 
left, rudimentary operculum on right. G, Worm showing extent of thoracic membrane. H-K, Bayonet collar chaetae. L, Thoracic uncini. 
M, Anterior abdominal uncini. N, Bundle of anterior abdominal flat trumpet chaetae. O, Tube of paratype, also showing worm with its two 
opercula in situ. P, Radioles with two well-developed opercula. Q, Two views of thorax showing extent of thoracic membranes. R-U, 
Bayonet collar chaetae. 






ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



85 




86 



T.G. PILLAI AND H.A. TEN HOVE 







Fig. 24 Spiraserpula iugoconvexa sp. nov. A-K, Paratypes. A, Dorsal view of erect tube part showing longitudinal external ridges, granular 
overlay, transverse ridges, triangular dorsal lobe of its aperture, and an unidentified transverse epibiont at its base. B, Ventral view of same 
tube showing the two small ventral lobes of the aperture, rounded ventral side and fine transverse growth ridges. C, Posterior coil of tube 
showing a damaged dorsal ridge. D, Radioles and operculum of the right side. E & F, Same operculum showing its zygomorphy and convex 
distal end. G & E, Dorsal and ventral views of thorax showing the collar and extent of the thoracic membranes. I, Juvenile paratype. J, 
Bayonet collar chaeta, lacking blade, but showing two teeth on the basal boss. K, Newly formed chaeta from within the fascicle. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



87 




Fig. 25 Spiraserpula iugoconvexa sp. nov. Holotype. A, Tube fragment showing irregular surface and peculiar structure with a semilunar 
opening (? Protulophila). B & C, Views of the opened tube showing the unserrated dorsal ridge, with a somewhat flatened ridge (C, 
bottom left). D, Tube fragment showing ventral ridge. E-G, Entire holotype, in three parts. E & F, Body showing the dorsal longitudinal 
groove, the apparently glandular ventral side of the abdomen and its peculiar outpouchings. G, Radioles with operculum on the left and 
rudimentary operculum on the right. G-J, Four views of the zygomorph operculum with convex distal end. K-N, Bayonet collar chaetae 
with long slender blade, short unserrated notch and two teeth (seemingly one tooth) on the basal boss. O, Thoracic uncini. P, Anterior 
abdominal uncini. Q, Bundle of anterior abdominal flat trumpet chaetae. 



88 



T.G. PILLAI AND HA. TEN HOVE 




Fig. 26 Spiraserpula snellii sp. nov. A, J-K, N-Q, V-W, Holotype. B-L, R-U, Paratypes. M, Juvenile. A & C, Tubes showing longitudinal 
pigment bands, transverse bands and thickenings (A). B, Tube fragment showing unserrated ventral ridge. E-H, tube fragments in their 
relative positions to the unopened tube (D) showing the unserrated ventral ridge which is thickened towards the middle of the tube (G,H). 
I, Tube fragment showing T-shaped ventral ridge. J-K, Holotype, showing operculum, radioles, and dorsal longitudinal groove along its 
body. L, Paratype, juvenile without operculum. M, Smaller juvenile worm, also without operculum. N-U, Bayonet collar chaetae. V & W, 
Thoracic uncini. X, Anterior abdominal uncini. 



Queensland (Australia): 2. Lizard Island, N of South Island, 
14.4°S 145. 3°E, reef front, sloping reef outside of lagoon and 
sandy bottom below, 10-17m, legit H. A. ten Hove, P. 
Hutchings and M. Reid, 5. hi. 1986, Stn.20 (3 specimens, 
ZMA V. Pol. 3734, AM W20342). 3. Lizard Island, Palfrey 



Island, S. of light-house; coral heads on sandy bottom, 7 m, 
legit H. A. ten Hove, 2. hi. 1986, Stn.17 (1 specimen in four 
fragments, BM(NH) 1992.65). 4. Lizard Island, S. South 
Island; sloping silty reef, little coral cover, legit H. A. ten 
Hove et al, Stn.21, 6. hi. 1986 (8+ specimens, ZMA V. Pol. 



ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



89 



3830, BM(NH) 1993.17, AM W21677). 5. Boulton Reef, on 
scleractinian coral (Thecopsammia regularis Gardiner 1899), 
USNM 78572, dry material, legit J. C. Lang, 31.vii.1973. H. 
Zibrowius, who identified this, kindly drew our attention to 
its serpulid epifauna. 

Loyalty Islands, E. of New Caledonia: 6. SW Pacific Lagoon 
of Beautemps-Beaupre Atoll; overhang 8 m, on heavily 
encrusted dendrophylliid scleractinian coral, scuba diving, 
dry material; MUSORSTOM 6 cruise, legit H. Zibrowius, 
17.ii.1989. 

Okinawa (Japan): 7. W. side of Sesoko Island, 2-3 m, on 
cliffside, in caves and grooves, scuba diving, on unidentified 
coral, legit S. Nakamura, 10.i.l989, dry material, USNM . H. 
Zibrowius kindly drew our attention to the serpulid epifauna. 
Sinai (Egypt): 8. Strait of Tiran, at Sharks Observatory, 
20-25 m; Nos. 210-213, legit H. A. ten Hove, 8.vi.l990 (2 
specimens, tubes, HUJ, ZMA V. Pol. 3886). 
Elat (Israel): 9. In front of Marine Biological Laboratory, 
20-25 m, coral rubble; Nos. 154, a-d, legit H. A. ten Hove, 
4.vi.l990. 10. Oil port, S. pier, 6-25 m, coral rubble and 
pillars of pier; Nos. 181,244, 311, 339, 340, legit H. A. ten 
Hove, 6.vi.l990 (3 specimens, several tubes, HUJ). 

Type locality. Taka Bone Rate (Flores Sea, Indonesia). 

Description. 

TUBES: Mustard coloured, with a pair of darker longitudinal 
bands in places along each flank, joined by transverse bands, 
especially just anterior to the thickenings found at intervals 
(Fig. 26, A,C). They may be coiled more or less parallel to 
one another in the horizontal plane, mutually bonded 
together or spread out on the substratum and branched in 
places. Their external diameter is quite small, only up to 
about 0.6 mm. Earlier formed portions of tubes may show 
narrow transverse wrinkles (Fig. 26, B,D). In fresh material 
the colour of the tube may be more brownish, and appears to 
fade to mustard after a few months in alcohol. 

ITS consist of an unserrated ventral ridge only (Fig. 26, 
B,E-I), which is T-shaped in cross-section towards its middle 
(Figs. 26, G, I; 3, F), and becomes progressively less thickened 
both anteriorly and posteriorly (Fig. 26, E,I). 
worms: The total length of the worms ranges from 2.2 mm in 
the case of a juvenile, to a little more that 12.3 mm in an 
older individual which lacks its radioles. The complete holo- 
type (Fig. 26, J) is only 5.8 mm long. The thoracic width in all 
the specimens is around 0.3 mm. 

An operculum may or may not be present. Younger 
specimens have radioles but lack opercula (Fig. 26, L,M); 
apparently opercula appear only in older worms (Fig. 26, 
A,J,K). The length of the operculum and peduncle in the 
holotype is 1.5 mm, the operculum 0.3 mm long and its 
diameter 0.2 mm. Its distal part is nearly globular 
(Fig.26,A,J,K) and, unlike the opercula of the other known 
members of the group, its margin is not divided into lobes, 
but shows about four pseudo-lobes, apparently caused by 
contraction in alcohol. Its proximal part is shaped like a 
narrow funnel, separated by a sharp constriction from the 
slender peduncle. A short filamentous rudimentary opercu- 
lum was observed in one specimen only. It appears likely 
that, like the operculum, they are developed in older worms. 
Pinnule-free tips of radioles short. Thoracic glands were not 
found. Some counts and meristic data are given in Table 20: 

The abdominal length in eight specimens ranged between 
11.2 and 1.0 mm, and the number of segments between 48 



Table 20 S. snellii sp. nov. Some meristic and other data of type 
series. 



No. of specimens (n=6) 
No. of radioles 


3 

5/5 


3 

4/4 








No. of specimens (n=8) 
No. of thoracic chaetal tufts 


2 
8/7 


1 
8/6 


2 

7/7 


2 
7/6 


1 

7/5 


No. of specimens (n=3) 
Thoracic membrane ends 


1 

4/4 


1 

4/3 


1 

3/3 







and 22, respectively, with capillaries on the last 6 or 7. 

Collar fascicles of older specimens bear about four fully 
formed bayonet chaetae and a developing one deep within. 
Each bayonet chaeta possesses a long serrated blade, a 
moderately long unserrated notch (1/3-1/4 the length of the 
entire blade), and several teeth on the basal boss (Fig. 26, 
N-U; PI. 5, F). Thoracic uncini (Fig. 26, V,W) and anterior 
abdominal uncini (Fig. 26, X; PI. 5, G) bear 4-6 and 4-5 teeth, 
respectively, in a single row. Flat trumpet-shaped chaetae are 
typical (PI. 5, H). 

Remarks. One single tube revealed 2 specimens: a parent 
with schizont closely appressed to its posterior end. Posteri- 
orly, the abdomen of the parent was abruptly tapering 
(dorso-ventrally), with long capillaries. Lying between those, 
the three pairs of radioles of the schizont could be found. It 
had a narrow, still not fully developed thorax with 7/6 chaetal 
tufts, followed by a well-formed abdomen with 17 chaetigers 
(the last 7 with capillaries). The entire schizont was folded 
over the ventral internal ridge. 

Collections from other localities. The specimens in 
sample 2 from Queensland agree with those in the Indonesian 
sample with regard to the overall mustard colour. Against 
this background colouration there are darker mustard to 
brown longitudinal bands, which are variable. One of the 
three available tubes has a pair of lateral longitudinal bands, 
lacking in places. The second tube has a thin median longitu- 
dinal stripe in addition. The third has a pair of mustard yellow 
longitudinal bands laterally, and a broad brownish median 
band which is partially divided into two bands by a narrow, 
yellow longitudinal band. 

They are coiled upon themselves either individually or 
mutually bonded together. The coils are more or less concen- 
tric, low, flattened against the substratum, and bonded 
together (Fig. 27, A). The maximum external tube diameter is 
1.2 mm. The granular overlay consists of a median longitudi- 
nal band made up of broad, transverse, forwardly-directed 
scutes, and a narrow band of smaller granules laterally 
(Fig. 27, A). At irregular intervals there are wavy, thickened, 
peristome-shaped transverse ridges. 

ITS agree with those of the Indonesian specimens. They 
consist of only an unserrated ventral ridge. Its edge is smooth 
and, in cross-sectional appearance, varies from being wedge- 
shaped to thickened and T-shaped at its maximal develop- 
ment (Fig. 27, B). The cross-bar of the T may also be curved 
outwards and bear a shallow longitudinal depression. The 
mid-ventral longitudinal abdominal groove is applied to this 
ridge. 

Three worms were removed from the tubes. One has a 
damaged thorax and an incomplete abdomen, while the 
second lacks the radioles of both sides, the third is broken in 4 
fragments. The former (Fig. 27, D), is 0.4 mm wide in the 



90 



T.G. PILLAI AND HA. TEN HOVE 



. 5mm 




ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



91 



thorax, and has 6 radioles and a short club-shaped rudimen- 
tary operculum on each side. The radioles are about 0.4 mm 
long and end in short pinnule-free tips. Two clusters of dark 
brown prostomial ocelli are present. There are 7 thoracic 
chaetal tufts on one side, but the number on the other side 
and the extent of the thoracic membranes cannot be deter- 
mined due to the damaged thorax. 

The first worm is spirally coiled along the substratal plane 
(Fig. 27, E). The length of the thorax and abdomen is 
13.8 mm, of which the posterior portion of about 1.18 mm is 
abruptly narrower than the rest of the abdomen. The entire 
length of the third worm is about 7.0 mm, radioles 1.3 mm, 
thorax and abdomen 5.7 mm; it has 7/6 radioles, and thoracic 
width of 0.35 mm. 

There are 7 thoracic chaetal tufts on each side in one worm, 
6/7 in the other. The thoracic membranes end between 
chaetal tufts 6 and 7 on the left, and 5 and 6 on the right 
(Fig. 27, E,F). Thoracic glands are absent. The number of 
abdominal segments is 102, with capillaries on the last 6. A 
mid-ventral longitudinal groove traverses the entire abdomen 
and thorax (Fig. 27, E, F). 

Each collar fascicle bears about 4 bayonet chaetae. They 
have long serrated blades, a short unserrated notch, and 
several teeth on the basal boss (Fig. 27, G-J). Thoracic uncini 
usually bear 4 teeth in a single row (Fig. 27, K). The anterior 
abdominal uncini also possess 4 teeth in a single row, but the 
3 posterior teeth are not closely appressed, as in the Indone- 
sian specimens (Fig. 27, L). Flat trumpet chaetae number 
about 3 per bundle. Since their edge is curved, details on the 
anterior tooth cannot be observed. 

The third sample from Lizard Island, Queensland agrees 
with the first and the Indonesian specimens in all important 
characters (Fig. 27, A-J). The fully formed operculum is an 
unlobed funnel with n shallow distal concavity (Fig. 28, C,D), 
while in the earlier stages it is spherical or nearly spherical 
(Fig.28, E-G). 

The dry tubes from Loyalty Islands (Fig.28, K-Q) 
appeared to be brownish, but regained the typical mustard 
colouration when immersed in alcohol. One fragment shows 
branching (Fig.28, K). Internally there is only a smooth 
ventral ridge (Fig.28, N-Q), which is clearly T-shaped in 
places (Fig.28, Q); it is markedly so and occupies a larger part 
of the lumen in some tubes which are comparatively very 
thick-walled (Fig.28, N). 

The collection from Ras Mohammed, Sinai, consists of 
tubes with fragments of worms (Fig.28, R-V). The tubes 
have an overall mustard colour, but the anterior portions 
(Fig.28, R,S) have pinkish peristomes, and a conspicuous 
granular overlay along the lateral borders of the attached 
portions (Fig.28, S). The medial overlay is scute-shaped, but 
not as prominent as in the first Lizard Island sample. A 
smooth ventral ridge is present (Fig.28, T-W), which is 
T-shaped in its fully formed condition (Fig.28, T,U,W). 
Those details of the worm that still could be observed (collar 
chaetae, ends of thoracic membranes) agree with the data 
given above. 

The samples from Elat, Israel, agree with regard to tube 



colouration, the smooth ventral longitudinal ridge and other 
important characters. No. 154 is a single specimen on a piece 
of coral rubble. The granular overlay is translucent in places; 
transverse scutes are not seen medially, but this may be 
because it is a juvenile. The worm has a total length of 
3.6 mm; reddish prostomial ocellar clusters are present; its 
thorax is 0.2 mm wide; gills 1.0 mm long, with short pinnule- 
free tips; the number of thoracic chaetal tufts L?/R7; its 
abdomen 1.9 mm long, with 24 segments and capillaries on 
the last 4. There are 3 bayonet chaetae per fascicle, each with 
an elongated blade, a short unserrated notch and several 
teeth on the basal boss. Anterior abdominal uncini bear 4 
teeth in a single row. An operculum had yet to be developed. 
However, an operculum was observed in sample 311. A 
schizont was separated from sample 244. 

Etymology. Named after the Indonesian-Dutch Snellius II 
Expedition which enabled the second author to collect exten- 
sively in Indonesian waters. 

Habitat and distribution. A reef dweller occurring at 
depths of about 15-30 m. Appears to be the most widely 
distributed species of the genus. Hitherto collected from the 
northern Red Sea, Indonesia (Flores Sea), Australia (Great 
Barrier Reef) and W. Pacific (S. Japan to New Caledonia). 

Spiraserpula lineatuba (Straughan, 1967) 

(Figs. 29, A-O; 30, A-M; 3, L; Pl.l, A, C & D, PI. 3, 

E-G) 

Synonymy. Serpula lineatuba Straughan, 1967, pp. 211-212, 
Fig.5a-g. 

Material examined. 

New South Wales: 1. Sydney, Long Reef, underside of rocks, 
LWS,27.ii.l965, legit D. Straughan (HOLOTYPE, 
AM4018). 2. Sydney, Long Reef, rocks just below LWS, 
Colloroy, Stn. 30, 27. i. 1964 (Topotypical material, 2 speci- 
mens and several tubes, AM4019, ZMA V. Pol. 3450, 
BM(NH) 1992.51). 3. Norah Head, at foot of light house, 
from bottom of tidal pools at low-tide, from undersides of 
boulders, legit H. A. ten Hove, 12. iv. 1986, Stn. 31 (5 out of 
several specimens, AM W20340). 4. Split Solitary Island, 
rocky island area with corals, algae and little sand, from 
ceiling of small cave, 12-19 m, legit H. A. ten Hove, P. 
Hutchings and R. Phipps, 26. iv. 1986, Stn. 36 (18 out of 
several specimens, ZMA V.Pol. 3709, USNM 130996, 
BM(NH) 1992.40-50, AM W20163, QM, NSMT). 5. South 
Solitary Island, S of light house, rocky area, cobbles and 
corals, little sand, 12-20 m, legit H. A. ten Hove, P. Hutch- 
ings and R. Phipps, 27. iv. 1986, Stn. 37 (3 out of several 
specimens, BM(NH) 1992.52-60). 

Type locality. Sydney, Long Reef (Australia). 

Description. 

According to the original description (Straughan, 1967), the 
tube is circular in cross-section, white, with a pair of dark 
pink lateral longitudinal stripes, pale pink dorsal surface. The 



Fig. 27 Spiraserpula snellii sp. nov. From Stn. 20, Lizard Island, Australia: A, Adult tube showing flattened coil form, granular overlay, 
which is scutate medially, granular laterally and has a transverse thickened peristome. B, Aggregation of tube fragments with unserrated 
ventral ridge, T-shaped in cross-section. C, Scutate juvenile tube with some transverse thickenings; granular overlay not yet developed. D, 
worm showing radioles, rudimentary operculum and collar. E, worm from tube figured in A, showing thoracic membrane, ventral 
longitudinal groove. F, Anterior part of latter, showing lack of apron and thoracic membrane ending on the 6th chaetiger on the left side. 
G-J, Bayonet chaetae, all from same fascicle. K, Thoracic uncini. L, Anterior abdominal uncini. 



92 



T.G. PILLAI AND HA. TEN HOVE 



total length of the worm ranges from 4.5-6.5 mm. There are 
4-6 pairs of branchiae with pinnule-free tips. A hollow 
operculum with about 22 lobes is present on one side, with a 
pseudoperculum on the other. The collar has a pair of lateral 
elongations on the median lobe. The thorax has 9 or 10 
segments on each side, and the bayonet collar chaetae have 2 
conical processes at the base of the blade. 

The holotype (AM W4018) is in very poor condition. When 
it was examined by the second author in 1979, the poorly 
preserved worm, still within its tube, lacked both an opercu- 
lum and a rudimentary operculum, although there appeared 
to have been one on one side and none on the other. Other 
observations were as follows: A cluster of pigmented ocelli 
present at the base of each branchial lobe; bayonet collar 
chaetae possess 2 conical teeth, with 1-3 accessory conical 
teeth; the anterior abdominal uncini of two types: some with 
a single row of teeth, others in which the posterior tooth is 
split into two; middle abdominal uncini appeared to possess 7 
simple teeth in side view; in edge view, however, four 
anterior teeth are single and the rest are rows of 3 minute 
teeth each. 

However, examination of topotypical material collected on 
the same date as the holotype and determined by Straughan 
(AM 4019, ZMA V.Pol. 3450) yielded the following addi- 
tional data: The tube has a pair of light pink longitudinal 
bands (Fig. 29, A), not clearly defined dark pink stripes as 
mentioned and figured in the original description. It is coiled, 
somewhat flattened against the substratum, but the free 
surface is rounded. The coils are bonded together. A granular 
overlay is present, but it is extremely fine and can only be 
seen in places, under special illumination (Fig. 29, F). A short 
erect portion is present (Fig. 29, A,B), with a four-lobed 
peristome similar to that of S. massiliensis. 

The most important data obtained during the present study 
of this topotypical material is that S. lineatuba has ITS. They 
consist of an unserrated dorsal ridge along the convex wall 
(Fig. 29, B,C,E,F), and a serrated ventral ridge along the 
opposite wall (Figs. 29, D,F; 3,L). The former may be high in 
the first formed coil (Fig. 29, E, F, bottom left), or low 
anteriorly (Fig. 29, B,C), and is wedge-, tongue- to somewhat 
T-shaped in cross-section. 

The worm is 6.5 mm long, its thorax is 0.5 mm wide, and 
its abdomen is 4.5 mm long. There are 5 radioles and a 
slender rudimentary operculum on one side. The median lobe 
of the collar has only one forwardly directed process, in 
contrast with the original description, indicating that this is a 
variable feature. There are 7 pairs of thoracic chaetal tufts, 
and the abdomen has 49 segments, with capillaries on the last 
19. Two clusters of prostomial ocelli are present and the 
thoracic membranes do not extend to the last thoracic chaeti- 
gers, but end on the fourth and fifth. 

There are 5 bayonet chaetae in each collar fascicle, each 
with a moderately long serrated blade, a moderately long 
unserrated notch which is 1/3-1/4 the length of the blade, and 
2 or 3 conical teeth on the basal boss (Fig. 29, G-J; PI. 3, E). 
In bayonets with two large teeth, a single accessory tooth may 



be present between them (Fig. 29, G,H,J). Thoracic uncini 
bear 5 or 6 teeth. As seen in edge view, in the outermost 
uncini of the row, 3 to 5 of the anterior teeth are single, while 
the remaining teeth are subdivided into 2 or more smaller 
teeth which form a short, rasp-shaped posterior cluster 
(Fig. 29, L). Anterior abdominal uncini are similar (Fig. 29, 
N). However, SEM of anterior abdominal uncini of another 
specimen showed a single row of teeth in edge view (PI. 3, F). 
It appears, therefore, that both types of uncini may some- 
times be present. Posterior abdominal uncini are rasp- 
shaped, except for the single anterior tooth. The uncini of the 
intermediate region show a transition between the two types. 
Flat trumpet chaetae number about 5-7 per fascicle (Fig. 29, 
N;P1.3,G). 

A more complete account of the species, however, was 
obtained from numerous well-preserved specimens collected 
in 1986 from Split Solitary Island. 

tubes: Have the colouration described above, including the 
pair of light to somewhat darker pink lateral longitudinal 
bands. They occur in aggregations of a few to numerous 
individuals, highly coiled amongst themselves and mutually 
bonded together, particularly at their bases (Fig. 29, O). 
Erect parts are sometimes present, and they may bear 
four-lobed peristomes (Fig. 30, A). The uncoiled part of one 
of the longest tubes measures 26.7 mm; together with its 
coiled part it is approximately 30.0 mm long, and its maxi- 
mum external width is 1.1 mm. 

ITS consist of an unserrated dorsal ridge, a serrated ventral 
ridge and, usually, a pair of accessory dorso-lateral ridges 
(Figs.29,0, middle left; 30, B,C; 3, L; Pl.l, A,C,D). The 
dorsal and ventral ridges of the tube are applied to corre- 
sponding longitudinal mid-dorsal and mid-ventral abdominal 
grooves (Figs. 30, D-F). 

Eighteen worms from Split Solitary Island provided impor- 
tant additional data. Measurements and other meristic data 
from 8 complete specimens of total lengths ranging between 
15.9 mm and 1.3 mm presented in Table 21 show that the 
worms can attain two and a half times the length mentioned 
in the original description. The maximum number of abdomi- 
nal segments counted is 89: 

Thirteen complete anterior ends all possess an operculum 
on one side, a rudimentary operculum on the other, and 5 
pairs of radioles. The pinnule-free tips are about 1/4 the 
length of the radioles and are as thick as the pinnules (Fig. 30, 
D,E). The length of the operculum together with its peduncle 
ranges between 0.8 mm in the smallest specimen to 1.6 mm 
in the largest; the length of the operculum itself from 0.3 mm 
to 0.7 mm, and its diameter from 0.4 mm to 0.6 mm, respec- 
tively. All the opercula are zygomorph (Fig. 30, D,F), their 
distal ends are concave and the radii end in somewhat pointed 
lobes. Many of the latter are actually double, the sub-dividing 
grooves being only about 1/3 the length of the main interra- 
dial grooves which extend to about half the opercular length. 
Thus the total number of about 17-23 radii end in about 
double the number of marginal lobes (Fig. 30, D-F). The 
constriction between the peduncle and the operculum is 



Fig. 28 Spiraserpula snellii sp. nov. A-J, from Stn. 21, Lizard Island, Australia. K-Q, From Loyalty Is. R-W, from Egypt. A-B, Tube 
lacking scutes and granular overlay, but with faint transverse grooves between transverse areas (representing scutes ?). D-E, Same worm 
with fully formed operculum. E-G, worms showing early vesicular operculum. H, Bayonet chaetae. I, Thoracic uncini. J, Anterior 
abdominal uncini. K, Anterior fragment of a tube showing branching and a peristome. L, Another fragment showing transverse ridges. M, 
Juvenile tube. N, Fractured end of a tube showing a thick wall and a T-shaped ventral ridge occupying most of its lumen. O-O, Tube 
fragments with varying form and thickness of the T-shaped ventral ridge. R & S, Anterior tube fragments, R with peristomes. T-W, tube 
aggregations with fractured ends showing the T-shaped ventral ridge. W, V, with longitudinal view of ventral ridge. 






ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 



93 




94 



T.G. PILLAI AND H.A. TEN HOVE 



Table 21 S. lineatuba (Straughan). Measurements and counts from 
Split Solitary Island material. 



Specimen 


Total 


Width of 


Length of 


No. of 


Capillaries 


no. 


length 


thorax 


abdomen 


abdominal 


on 






(mm) 


(mm) 


segments 




1 


15.9 


0.6 


13.2 


89 


13 


2 


15.7 


0.6 


12.6 


85 


13 


3 


15.7 


0.5 


12.7 


81 


12 


4 


13.9 


0.5 


10.9 


58 


17 


5 


13.0 


0.5 


12.7 


81 


13 


6 


10.1 


0.5 


10.9 


58 


17 


7 


9.3 


0.5 


6.8 


78 


16 


8 


1.3 


0.5 


0.4 


45 


25 



Table 22 S. lineatuba (Straughan). Number of thoracic chaetal 
tufts and extent of the thoracic membranes in specimens from 
Split Solitary Island. 



1 



1 



1 



No. of specimens (18) 1 
No. of thor. chaetal 
tufts 10/10 10/910/8 9/9 9/8 8/8 8/7 7/7 



No. of specimens 

(15) 
Thoracic membr. 

ends 



1 114 112 12 1 

7/5 6/4 5/5 5/4 5/3 4/4 4/3 4/2 3/3 3/2 



sharp, and the diameter of the distal end of the former varies 
from 1/3 to 2/3 that of the base of the latter. 

The median lobe of the collar shows one or more anteriorly 
directed processes in some specimens, none in others. Up to 8 
bayonet chaetae have been counted in a collar fascicle. A pair 
of ventral thoracic glands is present (Fig. 30, E). The number 
of thoracic chaetal tufts on each side varies from 7-10, and 
the thoracic membranes end on the 3rd to 6th chaetigers, as 
shown in Table 22. 

The specimens from South Solitary Island and Norah Head 
agree with the above description. 

Live material. No records. 

Habitat and distribution. The species occurs from the 
tidal zone down to about 20 m. It was very abundant on a 
ceiling of a small cave at a depth of 12-19 m, forming 
aggregations of up to 35 mm thick, and superficially 
resembles S. ypsilon from a similar habitat in the Cape Verde 
Islands. It has hitherto been collected only from N.S.W. 



Straughan, 29.iii.1986, legit H. A. ten Hove and P. Hutch- 
ings, Stn. 30 (HOLOTYPE, AM W20390). 

Type locality. Sydney, Long Reef (Australia). 

Description. 

tube: Pink, with whitish lateral attachment areas and very 
fine transverse wrinkles. The median tube parts are of a paler 
pink colour than the medio-lateral parts, in fresh material. A 
fine granular overlay is present, which can be seen at certain 
angles of illumination. The lateral borders of the tube are 
glassy and transparent. Irregularly laid along the outer sur- 
face of the tube, and more or less perpendicular to it, are 
small semilunar to crescentic discs (Fig. 31, A-C,E). They are 
very thin, pink, glassy and transparent, and their axes are at 
various angles to the longitudinal axis of the tube. Some of 
them are even attached to the substratum just outside the 
tube (Fig. 31, B). The maximum external diameter of the tube 
is 0.85 mm. 

ITS consist of a serrated ventral ridge along its concave 
wall (Fig. 31, E), and a smooth dorsal ridge. In addition, pink 
disks are found on the inside too, on either side of the 
serrated ventral ridge (Figs. 31, D; 3, M). In some cases the 
discs appear to be through and through the wall. The 
mid-ventral longitudinal groove of the abdomen (Fig. 31, F) is 
applied to the serrated ventral ridge. The worm appears to 
have a remarkable ability to adjust its abdominal segments in 
relation to these sharp discs within the tube. 
worm: Although only one specimen is available, it is com- 
plete (Fig. 31, F). Its total length is 7.7 mm, thoracic width 
0.56 mm; the abdomen is 6.6 mm long and has about 56 
segments, the last 20 with capillaries. There are 6 radioles and 
a rudimentary operculum on each side. A cluster of blackish 
prostomial ocelli is present at the base of the radioles on each 
side. There are 8 pairs of thoracic chaetal tufts. Where the 
thoracic membranes of the two sides end precisely is not clear 
since the thorax is highly contracted (Fig. 31, F,G), but an 
apron is absent. No thoracic glands were discernible. 

The number of bayonet chaetae, 6 in each collar fascicle, is 
high in relation to the size of the worm. Their serrated blades 
are moderately long, the unserrated notch is about 1/3 the 
length of the blade, and there are only 2-4 teeth on the basal 
boss (Fig.31,H-M). A few small accessory teeth may be 
present. Thoracic and anterior abdominal uncini bear about 6 
and 5 teeth, respectively, in a single row. 



Etymology, diskos (Gr.) = discus; pherein (Gr.) 
carry. 



to 



Spiraserpula discifera s